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Encyclopedia of Plant Viruses and Viroids [1st ed. 2019]
 978-81-322-3911-6, 978-81-322-3912-3

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K. Subramanya Sastry Bikash Mandal John Hammond S.W. Scott R.W. Briddon

Encyclopedia of Plant Viruses and Viroids

Encyclopedia of Plant Viruses and Viroids

K. Subramanya Sastry • Bikash Mandal John Hammond • S. W. Scott R. W. Briddon

Encyclopedia of Plant Viruses and Viroids

K. Subramanya Sastry Indian Council of Agricultural Research, IIHR Bengaluru, India

Bikash Mandal Indian Agricultural Research Institute New Delhi, India

Indian Council of Agricultural Research, IIOR and IIMR Hyderabad, India John Hammond USDA, Agricultural Research Service Beltsville, MD, USA

S. W. Scott Clemson University Clemson, SC, USA

R. W. Briddon John Innes Centre Norwich, UK

ISBN 978-81-322-3911-6 ISBN 978-81-322-3912-3 (eBook) ISBN 978-81-322-3913-0 (print and electronic bundle) https://doi.org/10.1007/978-81-322-3912-3 # Springer Nature India Private Limited 2019 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. This Springer imprint is published by the registered company Springer Nature India Private Limited The registered company address is: 7th Floor, Vijaya Building, 17 Barakhamba Road, New Delhi 110 001, India

DEDICATION

This book “Encyclopedia of Plant Viruses and Viroids” is dedicated to the International Committee on Taxonomy of Viruses (ICTV) (past and present executive, study groups and other members) for its continuous hard work providing systematic order in the taxonomy and nomenclature of viruses, including plant viruses and viroids We also dedicate this book to all the past and present research faculty and students of plant virology and allied subjects of the world for their rich research contributions and for advancing the study of plant viruses and viroids. Authors K. Subramanya Sastry Bikash Mandal John Hammond S. W. Scott R. W. Briddon

Foreword

It is a real pleasure to be invited to write a Foreword to this important and timely Encyclopedia as I have been involved in many similar earlier projects whose products are now sadly out of date: the CMI/AAB Descriptions of Plant Viruses, which Bryan Harrison and I started in 1970, and later the VIDE database, which morphed into the ICTVdB under Cornelia Büchen-Osmond. Furthermore, I am the sole survivor of the first committee of the International Committee on Taxonomy of Viruses (ICTV). It was elected at the Moscow Congress in 1966 and initially called the Provisional Committee on Nomenclature of Viruses. So I bring blessings from that august heavenly band! All of these earlier efforts by hundreds of virologists were the antecedents of this magnificent Encyclopedia. The careful collation and presentation of biological information of individual viruses and viroids is as important as ever for a variety of purposes. It is required for identifying novel pathogens and is also required for devising useful names for them. Note that virus and viroid species are man-made groupings, namely, “related strains/isolates, which are so similar that it is most convenient to know them by a single name.” Recent advances in molecular biology allow the genomic sequences of all subcellular pathogens to be determined routinely, and although numerical relationships can be calculated from those sequences, and used to form arbitrary categories, phenotypic information is indispensable for interpreting that information to form the most useful groupings to be given single names. This is clearly recognized by the 2018 ICTV Code of Nomenclature which includes the recently modernized Rule 3.20 and states “A species is a monophyletic group of viruses whose properties can be distinguished from those of other species by multiple criteria.” Arbitrary boundaries between taxa based on sequence similarity are not enough; biological (phenotypic) attributes are just as important. Up-to-date well-ordered information is also required by field pathologists, researchers, and, especially, those trying to devise control measures that minimize the economic damage caused by plant virus and viroid diseases. Appropriate integrated control programs must be based on accurate scientific information on the biology, mode of spread, etiology, and geography of pathogens. Finally, one must vii

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recognize the value of this Encyclopedia for the training of students, who are the scientists, pathologists, politicians, and administrators of the future. The plant viruses and viroids listed by the ICTV are only the tip of the iceberg, as the number of identified plant pathogens, including viruses and viroids, is growing at an alarming rate as they “emerge” from wild and local hosts and spread worldwide in the ever-increasing global trade in plants. Primary reports appear in the traditional journals and in metadata, like the CABI Crop Compendium, but a few are also recorded in such open-access databases as Wikipedia. Hence, there is a need for publications of multi-sourced and “digested information” like this Encyclopedia of Plant Viruses and Viroids, which has been assembled by one of India’s most experienced senior plant virologists, Prof. K. Subramanya Sastry, and a team of associated authors, Dr. Bikash Mandal, Dr. John Hammond, Prof. S.W. Scott, and Prof. R.W. Briddon. In this Encyclopedia, the authors have included around 1516 plant viruses and viroids, and the results of their Herculean endeavors are conveniently available both as hard copy and as an e-book. Canberra, Australia October 2019

Adrian Gibbs

Preface

The tropical, subtropical, and temperate environments harbor diverse plant pathogens. In almost all countries, a large number of plant pathogens cause substantial crop yield losses. Among them, plant viruses and viroids are significant plant pathogens that reduce plant vigor, yield, and quality of crop products. Since the discovery of tobacco mosaic virus at the end of the nineteenth century, numerous plant viruses and viroids have been discovered all over the world. At present, there are 1484 viruses and 32 viroids reported to affect plants. They are spread from plant to plant and from region to region by vectors, trade, and human activity relating to agriculture. As a result, over time some are distributed globally, whereas others have restricted distributions. Pioneer studies have demonstrated the complexity and diversity of viruses, and their interactions with vectors in relation to disease epidemics and crop losses. In many countries, researchers have made some headway in developing virus-resistant planting material and have also developed cultural, chemical, and integrated approaches for combating virus diseases. In addition, recently developed molecular techniques, ELISA, PCR, rolling circle amplification, next-generation sequencing, siRNA deep sequencing, and metagenomic approaches, are being utilized for accurate virus and viroid diagnosis / identification, with nucleotide sequence analyses perhaps being the most reliable means to identify the viruses. As an increasing number of viruses are being discovered, nomenclature and classification has become an increasingly challenging subject in virology. In the early days of virology, researchers named viruses based on disease symptoms and the name of the plant species from which they were initially isolated. This resulted in the accumulation of multiple names for one virus in the literature, causing a great deal of confusion. To oversee the rules and regulations of virus nomenclature and classification in all fields of virology, the International Committee on Taxonomy of Viruses (ICTV) was constituted 50 years ago. The formation of the ICTV was a unifying moment in the history of global research on viruses. The taxonomic information collated by the ICTV is published in the form of a consolidated report at about 5-year intervals. The first report was published in 1971 and the tenth report in 2017. In the initial reports, plant viruses were divided into groups unlike viruses infecting vertebrates or bacteria. The proposal and acceptance of the species concept for viruses was an important milestone in virus classification that led to the formation of taxonomic structure for viruses as orders, families, genera, and species. The virus species concept was originally adopted in 1991 by the ICTV and fully implemented in the seventh report published in 1999 which marks the beginning of the modern era of virus taxonomy. In many instances, definite association of a specific virus with a disease is difficult to achieve due either to the induction of similar symptoms by different viruses or to the presence of mixed infections of related and unrelated viruses. The “Descriptions of Plant Viruses” were originally published by the Commonwealth Mycological Institute (CMI) and the Association of Applied Biologists (AAB) and then by the Virus Identification Data Exchange (VIDE), International Committee on Taxonomy of ix

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Preface

Viruses Virus Database, and CAB International, and these publications included viruses described up to 1996. Subsequently, a large number of viruses and viroids have been identified, and the taxonomic positions of many previously known viruses have been revised. Currently, the ICTV provides the list of approved virus species but not their full description. However, a full description of ecological, physical, biological, and molecular properties is essential for the identification of each virus. To develop a comprehensive description of the global plant viruses and viroids, we have covered the published literature on plant virology up to 2018. Sustained efforts over 5 years have resulted in the synthesis of this book, Encyclopedia of Plant Viruses and Viroids. It covers the description of 1516 plant viruses and viroids. Each of the viruses and viroids is described to the extent possible based on the available data, including taxonomic position, geographic distribution, symptoms, hosts, transmission, virion morphology, and genomic properties. Synonyms by which viruses have been reported previously are also included to aid in the interpretation of older reports in the literature. The unique aspect of this encyclopedia is that all viruses and viroids known to infect a plant species globally are described and the plant species are arranged in alphabetical order of the scientific name of the plant. Genomic features and the relationships of each virus are included in the description, which is one of the most important criteria for virus or viroid species identification. It must be pointed out that although the widespread use of nextgeneration sequencing to identify and classify viruses is rapidly adding to the number of reported plant viruses, the information available for these newly described viruses (the host range, symptomatology, distribution, incidence, vector relationships, impact on plant yields, etc.) is limited. We hope that this encyclopedia will be of great help by serving as a ready-reference source for background information on host plants and viruses/viroids for researchers and academics. November 2019

K. Subramanya Sastry Bikash Mandal John Hammond S. W. Scott R. W. Briddon Authors

Acknowledgments

For the finalization of this reference book entitled Encyclopedia of Plant Viruses and Viroids, the authors have benefited immensely from critical and constructive suggestions made by a large number of plant virologists in both national and international scientific communities, viz., Professors M. J. Adams, M. H. V. Van Regenmortel, A. J. Gibbs, T. A. Zitter, the late G. Loebenstein, H. R. Pappu, Anupam Varma, P. Lavakumar, V. Muniyappa, M. Krishna Reddy, F. J. Morales, T. Sano, G. P. Rao, D. V. R. Saigopal, P. Sreenivasulu, P. Parvatha Reddy, B. L. Subba Rao, Kokkarachedu Sridhar, G. Nagaraja, R. A. Naidu, the late Karl Maramorosch, the late J. M. Thresh, and the late M. V. Nayudu, and from other scholarly academicians, friends, and colleagues who provided their timely guidance and assistance during the finalization of this reference book, and the authors are highly grateful to them. We are also highly indebted to all of those contributing their expertise to the International Committee on Taxonomy of Viruses (ICTV) for their dedicated work in preparing the tenth report on Virus Taxonomy (2018), which we have utilized as the basis for plant virus and viroid classification in this book. The authors express their indebtedness to Prof. Andrew J. Davison (President); Prof. Elliot J. Lefkowitz (Editor-in-Chief ICTV Report); study group members and all other members of ICTV for facilitating us to use partial information / descriptions of plant virus families, genus and species provided in the tenth ICTVonline while finalizing this reference book. If there are any inadvertent discrepancies between the taxonomic classifications shown here and the ICTV tenth report, the authors defer to the ICTV. The authors and the Springer publishers are highly thankful to all the administrative and scientific authorities of international organizations like the FAO, EPPO, APS, and CABI and various publishers for their prompt help in providing information and permissions. We express our sincere gratitude to Mr. C. Nagaraja for computerizing the book and for the secretarial assistance throughout the preparation of this reference book. We are also grateful to Miss. B. Pavithra of SPi Global, Pondicherry, for taking keen interest during the finalization of proofs. Prof. K. Subramanya Sastry expresses his gratitude to his wife Mrs. B. N. K. Kumari for her continuous support during the preparation of this book and also to his parents the late Kokkarachedu Panduranga Sastry and the late K. Subhadramma for their eternal blessings. We thank Mokkapati Muralidhar, Mokkapati Padmavathi, and K. Sridhar for their co-operation during the completion of this book. Dr. John Hammond gratefully recognizes his wife, Dr. Rosemarie Hammond, for her sacrifices and support over the period of preparation of this Encyclopedia, and his colleagues, in particular Drs. Dimitre Mollov and Ramon Jordan, for the assistance with some sections. Dr. Bikash Mandal is highly thankful to his better half, Dr. Seema Mandal for her encouragement and support during the preparation of the Encyclopaedia. We thank Elsevier, CABI, Springer, Academic Press, and other publishers for providing permissions for using some of the information from their earlier publications. The authors gratefully appreciate and acknowledge the staff of Springer publishers particularly Dr. Mamta Kapila, Zuzana Bernhart, Tina

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Acknowledgments

Shelton and Pavithra Balakrishnan for the continuous support and the meticulous care taken in bringing out this publication at an early date. We hope that this book will be of value and interest to many plant pathology researchers particularly plant virologists, teachers, students of agriculture, seed companies, and researchers at quarantine stations, to serve as a comprehensive and accurate reference book for the plant virus and viroid diseases of all crops with some of the latest information on classification, transmission, genomic characters, and other aspects. We shall deem it an honor and reward if readers find this book is of great help by serving as a readyreference source to obtain the background and latest information on the host plants and viruses/viroids. We welcome the submission of suggestions and comments for the improvement of this Encyclopedia/ reference book in the future editions to e-mail: [email protected]. K. Subramanya Sastry Bikash Mandal John Hammond S. W. Scott R. W. Briddon Authors

Introduction

The Overview of Description of Plant Viruses and Viroids Virology, a branch of biological science, developed after the discovery of tobacco mosaic virus in 1898. Viruses are known to infect all sorts of living organisms, and those with host ranges known to be restricted within the plant kingdom are referred to as plant viruses. Although a few plant viruses are known to also infect their insect vectors, so far none of the plant viruses are known to infect higher animals including human beings. Plant viruses range in sizes from 16 to 2000 nm and are generally organized with nucleic acids and protein in either cubic or helical symmetry resulting in an isometric or rod-shaped architecture. A second class of infectious agents previously presumed to be viruses, and typified by the agent of potato spindle tuber disease, was shown to consist of only a covalently closed circular strand of single-stranded ribonucleic acid, lacking any protein coat; these agents were referred to as viroids (Diener 1971) and were later shown to lack any protein-coding capacity (e.g., Flores et al. 1997). Plant viruses are an important constraint on world agriculture. Of all the phytopathogens, studies of plant viruses have received special attention as they cause significant losses to crop yield and quality, and the management of the diseases requires a knowledge of both the associated virus and any vector involved in its transmission. Numerous plant viruses are presently known globally, and almost all crop species are affected by one or more viruses. Over the last three decades, the application of molecular techniques has revolutionized our understanding of plant viruses. Viruses are simple biological entities that undergo continuous and rapid changes in their genetic makeup and thereby emerge as numerous variants that pose a challenge for their identification and classification. No single property of viruses is adequate to identify them. A set of characters such as the disease they incite, the host plants they infect, the mode of transmission, specific vector, virion morphology, and more recently genome sequence and organization are used to describe a virus. The genome sequence is the ultimate feature in virus identification, which establishes genetic relationships and the evolutionary history of a virus. Therefore sequence information has become of paramount importance in viral taxonomy. However, genomic characteristics do not by themselves justify taxonomic allocations, and the wish to record phylogeny should not overshadow the importance of the other phenotypic and biological properties. The definition of a species as a polythetic class was endorsed by the ICTV, and species became the lowest level in virus classification; unfortunately, many virologists (Pringle 1991) thought that this definition would provide them with guidelines for the establishment and demarcation of new virus species and allow decisions as to whether a virus was a member of a particular species. This led to a continuing debate about the presumed usefulness of a species definition for creating new species taxa and identifying their members.

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Introduction

Frequently, the disease caused by a virus is difficult to understand due to the symptoms resulting from a mixed infection of viruses that may or may not be related. The reverse genetic approach to develop infectious clones of viruses provides an opportunity to determine the authentic disease symptoms caused by a virus. Epidemiology, biology, molecular biology, and interactions with vector and host are important areas of studies in plant virology with a goal of offering a solution to the problems imposed by plant viruses in sustainable agricultural production. The intensification of crop cultivation, environmental change, and international trade in agricultural products has resulted in more viral disease problems across the world. The application of molecular tools and techniques and genome sequence information has identified numerous new plant viruses. Over time, nomenclature and classification of viruses have been emerged as a complex and intense area of study in virology.

Taxonomy of Plant Viruses Plant viruses were initially identified based on the disease symptoms and the name of the host from which the virus was first isolated, e.g., tobacco mosaic virus. However, with the discovery of greater numbers of plant viruses, some of the recommended criteria were inadequate for classification. Further, more difficulties were encountered when it was shown that a particular virus could cause different symptoms in different plant species and different viruses could cause the same symptoms in a single plant species and has led to the misidentification of plant viruses. The earliest classification of plant viruses was published by James Johnson in 1927. He used a system to differentiate the viruses by naming them using the host name where the disease was first observed, the word “virus,” and a number (e.g., tobacco virus 1 for TMV). Other efforts for grouping plant viruses were made during this period by Smith (1937) and Holmes (1939) with minor modifications of the system published by Johnson (1927). Johnson and Hoggan (1935) further revised and developed a classification system using five characters: symptom, host, transmission, longevity in vitro, and thermal inactivation point. In the period from 1915 to 1940, several insect vectors, including aphids, whiteflies, thrips, planthoppers, and leafhoppers, were identified as being able to transmit plant viruses with extraordinary specificity, and this property was used as an important criterion for differentiation of viral disease from diseases caused by fungi and in the taxonomy of plant viruses. Fungi and nematodes were later discovered to be vectors of some plant viruses. During the years from 1935 to 1960, new technologies, such as electron microscopy, centrifugation, electrophoresis, and methodologies such as negative staining, virus purification, and crystallography were used to determine the physical and biochemical properties of viruses. Virion morphology, capsid protein symmetry, the presence or absence of an envelope, physical properties (buoyant density, molecular mass, sedimentation coefficient, virion stability), and chemical properties (type of nucleic acid [RNA or DNA], strandedness [single or double], topology [linear or circular], polarity [positive or negative] have been used as relevant physicochemical characters in the taxonomy of viruses. Another system proposed for virus classification was the use of cryptograms as an addition to the vernacular (common) name of the virus (Gibbs et al. 1966; Gibbs and Harrison 1968; Gibbs 1968). The cryptogram consisted of four pairs of symbols, which represented (i) the type, and strandedness of the genomic nucleic acid; (ii) the molecular weight of the nucleic acid, and the percentage of nucleic acid in the viral particle; (iii) the outline of the particle, and of the nucleocapsid; and (iv) the types of host, and types of vector. For example, the cryptogram of the Potexvirus group, as it was then described, is [R/1: 2.2/ 6: E/E: S/0] (RNA genome/single stranded: 2.2  106 MW/6% nucleic acid): elongated particle/ nucleocapsid: seed plant hosts/spreads without a vector. The cryptogram system drew attention to the level of data required for adequate classification, but was not fully available for many known viruses; however, the usage was restricted almost entirely to the plant virus community, with negligible adoption

Introduction

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by animal or bacterial virologists, despite the potential for applicability. Furthermore, the cryptogram system was not readily expanded to allow for the added complexities of the multipartite viruses then being identified, or for the variety of replication strategies including reverse transcription, or negative and ambisense genomes. The cryptogram system was never officially adopted by the ICTV but was nonetheless used in ICTV reports for a few years, before being discontinued for any ICTV use after 1977. Before any plant viral nucleotide sequences were determined, an important milestone was reached with the determination of the coat protein amino acid sequence of TMV (Tsugita et al. 1960). Protein sequencing was used to examine the phylogeny of many other organisms, establishing presumptive evolutionary trees using highly conserved proteins (e.g., Dayhoff et al. 1978), and many of the methods used for comparing nucleotide sequences were derived from those developed for analysis of amino acid sequences. The genomic sequence of a virus is considered the most important criterion in the classification of viruses. However, there is no reason to assume that when virus species are demarcated only on the basis of genome sequences and a derived hypothetical phylogeny, this will necessarily produce a classification that is more correct, relevant, or useful than a classification based on all the phenotypic properties of a virus (Calisher et al. 1995; Mahner and Bunge 1997). The generation of plant virus genome sequence, however, provides critical information for the identification and differentiation of plant viruses. Viral genome sequence was very useful in resolving the confusion over the synonymous identity of several plant viruses in the ninth report of the International Committee on Taxonomy of Viruses (ICTV) where the definite plant virus species as well as tentative members of a genus were listed (King et al. 2012). During the formative phase of virus taxonomy, various systems or approaches were articulated for virus nomenclature and classification, but none was adopted uniformly. It was increasingly felt that a uniform and internationally agreed system of nomenclature and classification was necessary to accommodate known and as yet unidentified viruses. In order to develop internationally agreed rules and regulations for nomenclature and classification of viruses, the “International Committee on Nomenclature of Viruses” (ICNV) was established during the ninth congress of the “International Association of Microbiological Societies” held in Moscow in 1966. Subsequently, the ICNV changed its name to ICTV in 1975 and became a part of the “Virology Division” of the “International Union of Microbiological Societies” (IUMS). In addition to developing a universal taxonomy for all viruses, viroids, and satellite viruses, the ICTV also performs an important role in communicating taxonomic decisions and maintenance of an upto-date index of virus species (Van Regenmortel 1990; Van Regenmortel et al. 1997, 2000a, b). The organizational structure of the ICTV is comprised of the executive committee, subcommittees, and study groups. New taxonomic proposals are scrutinized rigorously by members of the study group, subcommittee, and executive committee and finally accepted following ratification by a vote of the members of the various subcommittees, national members, and life members. The ICTV report provides taxonomic standards and official recognition of new viruses and approves creation of virus taxa. Every 5 years ICTV study groups revise the status of viruses and publish a consolidated report of taxonomy of viruses. So far the ICTV has published the virus taxonomy up to the ninth report in the form of a hard copy book in 1971, 1976, 1979, 1982, 1991, 1995, 2000, 2005, and 2012 (Wildy 1971; Fenner 1976; Matthews 1979, 1982; Francki et al. 1991; Murphy et al. 1995; Van Regenmortel et al. 2000a, b; Fauquet et al. 2005; King et al. 2012). It has now been decided by the ICTV that the taxonomic report will no longer be published in the form of a hard copy book and the 2017 classification report will be available online as an open access resource at http://www.ictvonline.org/report (Adams et al. 2017). In the ninth report of the ICTV, 23 families and 114 genera and 1286 plant viruses were included. Even 30 viroids infecting plants are grouped by ninth ICTV report into 8 genera and 2 families (King et al. 2012). In the tenth ICTV report (2018b.v1), among the total viruses, the plant viruses are in 26 families, 118 genera, and 1516 plant virus species. Even 32 viroids infecting plants are grouped in the tenth ICTV report into 8 genera and 2 families (Anon 2017). The current ICTV taxonomy can be found at talk. ictvonline.org/taxonomy.

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An important turning point in the development of plant virology was in 1960, when the amino acid sequence of the capsid protein of TMV was determined (Tsugita et al. 1960). Over the subsequent decades, viruses have been studied in greater detail, and the complete genome sequence of even the largest virus is now routinely determined. Each genome sequence provides the complete genetic character of an individual virus isolate. Much of the phenotypic information encoded by these sequences can be determined. Comparisons of the gene sequences and the proteins they encode provide estimates of phylogenetic relatedness and past evolutionary history, which are important types of information required in classification of viruses. Metagenomic data are changing our views on virus diversity and are therefore challenging the way in which we recognize and classify viruses (Simmonds 2015; Van Regenmortel 2016b). The viruses detected using metagenomic sequencing data can be described in three different ways: (1) known knowns, virus species or isolates that are already known to be in the environment being surveyed; (2) unknown knowns, new virus species or isolates of a known family or known viruses that have not been found previously in the surveyed environment; and (3) unknown unknowns, viruses that are completely novel and share little to no sequence similarity with other known viruses. Sequencing data for each instance can be analyzed differently based on the questions being addressed. The removal of non-viral sequences from the sample either before or after sequencing will, of course, increase the chances of identifying viruses within a metagenomic sequence dataset, so care should be taken in both sample preparation before sequencing and manipulation of sequence data after sequencing. For known knowns and unknown knowns, the screening of the sequence dataset for the presence of known viruses can drastically reduce the amount of time needed for analysis and as such, aid detection and identification of viruses (Stobbe et al. 2013). Metagenomics offers us a unique tool to elucidate the current state of viruses in plants and the roles different viruses play in virus: virus and virus: host interactions. It has been proposed that it should be possible to incorporate viral metagenomic sequences in the existing official virus classification system (Van Regenmortel 2016c). An important milestone of virus taxonomy is the recognition and adoption of the virus species concept. Van Regenmortel (1989) defined virus species as “A viral species is a polythetic class of viruses that constitutes a replicating lineage and occupies a particular ecological niche,” which was accepted by the ICTV in 1991 and fully adopted in the seventh report published in 1999. Virus classification deals with the abstract classes of viruses. A virus is an entity with distinct and independent existence, and it should not be confused with the virus species that is a conceptual construction of human minds and not real. Brandes and Wetter (1959), following the 1950 recommendation of the Virus Subcommittee of the International Nomenclature Committee for the first-listed and most important characteristic of the virus particles, “morphology and methods of reproduction,” proposed a classification of rod- and flexuousshaped plant viruses on the basis of morphological differences. They subdivided various viruses into 12 classes based on particle length and rigidity/flexibility and grouped these into 4 groups each containing 3 of the original 12, namely, (i) rigid rods of ~130 nm (barley stripe mosaic [hordei]virus and soil-borne wheat mosaic [furo]virus), ~180 nm (potato stem mottle virus = tobacco rattle [tobra] virus), and ~300 nm (tobacco mosaic virus and three other tobamoviruses); (ii) flexible filaments of ~480 nm (white clover mosaic virus), ~515 nm (potato virus X, cactus virus 1 = cactus virus X), an intermediate group with reported size overlapping the ~480 and ~515 nm classes (cymbidium mosaic virus), and ~580 nm (potato aucuba mosaic virus), all now recognized as potexviruses; (iii) rigid to slightly flexible particles of ~620 (Wisconsin pea streak virus) or ~650 nm (red clover vein mosaic virus, carnation latent virus, potato virus S, potato virus M, and cactus virus 2) transmitted by aphids, all currently definitive or tentative carlaviruses, and ~700 nm (wheat streak mosaic virus) transmitted by mites, now classified as a tritimovirus; and (iv) flexible threads of ~730 nm (beet mosaic virus, potato virus A, potato virus Y, tobacco etch virus, and henbane mosaic virus – all potyviruses), ~750 nm (bean

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common mosaic virus, bean yellow mosaic virus, pea mosaic virus, soybean mosaic virus, turnip mosaic virus, cocksfoot [streak] mosaic virus, lettuce mosaic virus, and sorghum red stripe virus [= sugarcane mosaic virus] – also all potyviruses), and ~1250 nm (beet yellows virus – now recognized as a closterovirus). By the time of the first report of the ICTV (1971), potato virus X was recognized as the type species of the Potato virus X group, potato virus Y as type member of the Potato virus Y group, and carnation latent virus as a member of the Carnation latent virus group; in the second report (1976), these were, respectively, the Potexvirus group, Potyvirus group, and Carlavirus group; in 1993 the “groups” were elevated to genus status, with the genus Potyvirus also classified within the family Potyviridae. The genera Potexvirus and Carlavirus were included in the family Flexiviridae in 2004 and divided in 2009 between the Alphaflexiviridae and Betaflexiviridae, respectively, of the order Tymovirales. Wheat streak mosaic virus was initially classified by the ICTV as a member of the genus Rymovirus, family Potyviridae, but transferred to the new genus Tritimovirus (still Potyviridae) in 1998. These examples show the general stability of plant virus classification throughout the history of the ICTV, with addition of further levels of discrimination as more information becomes available; most of the new information leading to updated taxonomic placement has come directly from new sequence information and phylogeny, and several previously recognized species have either been folded into other taxa as synonyms have been identified, or some species previously established only on biological information and lacking sequence information have been abolished as recognized species. The latest plant virus and viroid classification is provided in Table 1.

Nomenclature of Viruses In deciding how to write a virus name, it is important to know whether it is the name of a virus taxon (a species, genus, subfamily, family, or order) or the name of a virus as a physical or genetic entity replicating in a host. Virologists have encountered problems in naming newly discovered viruses that differ by genome organization or sequence identity from those which induce similar disease symptoms. This situation is frequently encountered while naming members of the genus of the family Geminiviridae. The ICTV provides a stable and uniform nomenclature system of the virus species, genus, family, and order. The orthography rules are available at https://talk.ictvonline.org/information/ w/faq/386. The ICTV-recognized taxa (order, family, genus, and species) are written in italic with the first letter capitalized. A virus name should never be italicized, even when it includes the name of a host species or genus, and should be written in lower case. This ensures that it is distinguishable from a species name, which otherwise might be identical. The first letters of words in a virus name, including the first word, should only begin with a capital when these words are proper nouns (including host genus names but not virus genus names) or start a sentence. Single letters in virus names, including alphanumerical strain designations, may be capitalized. In most texts, virus names are used much more frequently than species names and may, therefore, be abbreviated. In the case of virus strain and isolate, the name is neither italicized nor is the first letter capitalized, unless it is a proper noun or part of the scientific name. The abbreviated version of the virus name is used in the form of a unique acronym (or almost unique; there are a few exceptions) for each virus isolate or strain. However, as a species includes a group of close variants (isolates/strain), the species name is never abbreviated. In a manuscript, often virus family, genus, and species names are used in singular and plural form without referring to the taxonomic terms. Under such informal usage, vernacular terms are used where they are neither italicized nor have the first letter capitalized, e.g., begomoviruses and isolates of tomato leaf curl New Delhi virus (Van Regenmortel and Fauquet 2002).

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Table 1 Order, family, genus and type species of plant viruses and viroids classification (Source: ICTV 2018b.v1) I. Plant viruses Nature of genome (+) sense ssRNA Viruses

Order Picornavirales

Family Secoviridae

Tymovirales

Alphaflexiviridae

Tymovirales

Betaflexiviridae

Genus Comovirus Fabavirus Nepovirus Cheravirus Sadwavirus Sequivirus Torradovirus Waikavirus Unassigned Unassigned Unassigned Unassigned Unassigned Allexivirus Lolavirus Mandarivirus Platypuvirus Potexvirus Carlavirus Foveavirus Robigovirus Unassigned Unassigned Unassigned Capillovirus Chordovirus Citrivirus Divavirus Prunevirus

Tymovirales

Tymoviridae

Unassigned Unassigned Unassigned

Benyviridae Botourmiaviridae Bromoviridae

Tepovirus Trichovirus Vitivirus Wamavirus Maculavirus Marafivirus Tymovirus Unassigned Benyvirus Ourmiavirus Alfamovirus Anulavirus Bromovirus Cucumovirus Ilarvirus Oleavirus

Type species Cowpea mosaic virus Broad bean wilt virus 1 Tobacco ringspot virus Cherry rasp leaf virus Satsuma dwarf virus Parsnip yellow fleck virus Tomato torrado virus Rice tungro spherical virus Black raspberry necrosis virus Chocolate lily virus A Dioscorea mosaic associated virus Strawberry latent ringspot virus Strawberry mottle virus Shallot virus X Lolium latent virus Indian citrus ringspot virus Donkey orchid symptomless virus Potato virus X Carnation latent virus Apple stem pitting virus Cherry necrotic rusty mottle virus Banana mild mosaic virus Banana virus X Sugarcane striate mosaicassociated virus Apple stem grooving virus Carrot Ch virus 1 Citrus leaf blotch virus Diuris virus A Apricot vein clearing associated virus Potato virus T Apple chlorotic leaf spot virus Grapevine virus A Watermelon virus A Grapevine fleck virus Maize rayado fino virus Turnip yellow mosaic virus Poinsettia mosaic virus Beet necrotic yellow vein virus Ourmia melon virus Alfalfa mosaic virus Pelargonium zonate spot virus Brome mosaic virus Cucumber mosaic virus Tobacco streak virus Olive latent virus 2 (continued)

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xix

Table 1 (continued) I. Plant viruses Nature of genome

Order Unassigned

Family Closteroviridae

Genus Ampelovirus Closterovirus Crinivirus Velarivirus Unassigned Unassigned Unassigned Unassigned Unassigned Unassigned

Unassigned

Unassigned

Kitaviridae

Luteoviridae

Unassigned

Potyviridae

Unassigned

Solemoviridae

Unassigned

Tombusviridae

Unassigned Blunervirus Cilevirus Higrevirus Enamovirus Luteovirus Polerovirus Unassigned Unassigned Unassigned Unassigned Unassigned Unassigned Unassigned Bevemovirus Brambyvirus Bymovirus Ipomovirus Macluravirus Poacevirus Potyvirus Roymovirus Rymovirus Tritimovirus Unassigned Unassigned Unassigned Polemovirus Sobemovirus Umbravirus Alphacarmovirus Alphanecrovirus

Type species Grapevine leafroll-associated virus 3 Beet yellows virus Lettuce infectious yellows virus Grapevine leafroll-associated virus 7 Actinidia virus 1 Alligatorweed stunting virus Blueberry virus A Megakepasma mosaic virus Mint vein banding-associated virus Olive leaf yellowing-associated virus Persimmon virus B Blueberry necrotic ring blotch virus Citrus leprosis virus C Hibiscus green spot virus 2 Pea enation mosaic virus 1 Barley yellow dwarf virus PAV Potato leafroll virus Barley yellow dwarf virus GPV Barley yellow dwarf virus SGV Chickpea stunt disease associated virus Groundnut rosette assistor virus Indonesian soybean dwarf virus Sweet potato leaf speckling virus Tobacco necrotic dwarf virus Bellflower veinal mottle virus Blackberry virus Y Barley yellow mosaic virus Sweet potato mild mottle virus Maclura mosaic virus Triticum mosaic virus Potato virus Y Rose yellow mosaic virus Ryegrass mosaic virus Wheat streak mosaic virus Common reed chlorotic stripe virus Longan witches broom-associated virus Spartina mottle virus Poinsettia latent virus Southern bean mosaic virus Carrot mottle virus Carnation mottle virus Tobacco necrosis virus A (continued)

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Introduction

Table 1 (continued) I. Plant viruses Nature of genome

(-) sense ssRNA Viruses

Order

Family

Unassigned

Virgaviridae

Unassigned Serpentovirales Mononegavirales

Unassigned Aspiviridae Rhabdoviridae

Genus Aureusvirus Betacarmovirus Betanecrovirus Gallantivirus Gammacarmovirus Macanavirus Machlomovirus Panicovirus Pelarspovirus Tombusvirus Zeavirus Unassigned Unassigned Unassigned Unassigned Dianthovirus Avenavirus Furovirus Goravirus Hordeivirus Pecluvirus Pomovirus Tobamovirus Tobravirus Idaeovirus Ophiovirus Cytorhabdovirus Dichorhavirus Nucleorhabdovirus Varicosavirus

(+/-) sense ssRNA viruses dsRNA viruses

Bunyavirales

Fimoviridae

Emaravirus

Bunyavirales Bunyavirales Bunyavirales

Phenuiviridae Unassigned Tospoviridae

Tenuivirus Coguvirus Orthotospovirus

Unassigned Unassigned Unassigned

Amalgaviridae Endornaviridae Partitiviridae

Amalgavirus Alphaendornavirus Alphapartitivirus Betapartitivirus Deltapartitivirus Unassigned Unassigned Unassigned

Type species Pothos latent virus Turnip crinkle virus Tobacco necrosis virus D Galinsoga mosaic virus Melon necrotic spot virus Furcraea necrotic streak virus Maize chlorotic mottle virus Panicum mosaic virus Pelargonium line pattern virus Tomato bushy stunt virus Maize necrotic streak virus Bean mild mosaic virus Chenopodium necrosis virus Cucumber soil-borne virus Trailing lespedeza virus 1 Carnation ringspot virus Oat chlorotic stunt virus Soil-borne wheat mosaic virus Gentian ovary ringspot virus Barley stripe mosaic virus Peanut clump virus Potato mop-top virus Tobacco mosaic virus Tobacco rattle virus Raspberry bushy dwarf virus Citrus psorosis ophiovirus Lettuce necrotic yellows cytorhabdovirus Orchid fleck dichorhavirus Potato yellow dwarf nucleorhabdovirus Lettuce big-vein associated varicosavirus European mountain ash ringspotassociated emaravirus Rice stripe tenuivirus Citrus coguvirus Tomato spotted wilt tospovirus

Southern tomato virus Oryza sativa alphaendornavirus White clover cryptic virus 1 Atkinsonella hypoxylon virus Pepper cryptic virus 1 Alfalfa cryptic virus 1 Carnation cryptic virus 1 Carrot temperate virus 1 (continued)

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xxi

Table 1 (continued) I. Plant viruses Nature of genome

Order

Family

Unassigned

Reoviridae

Caulimoviridae

Badnavirus Caulimovirus Cavemovirus Petuvirus Rosadnavirus Solendovirus Soymovirus Tungrovirus

Type species Carrot temperate virus 2 Carrot temperate virus 3 Carrot temperate virus 4 Hop trefoil cryptic virus 1 Hop trefoil cryptic virus 3 Radish yellow edge virus Ryegrass cryptic virus Spinach temperate virus White clover cryptic virus 3 Wound tumor virus Fiji disease virus Rice ragged stunt virus Banana bunchy top virus Subterranean clover stunt virus Coconut foliar decay virus Beet curly top Iran virus Bean golden yellow mosaic virus Euphorbia caput-medusae latent virus Beet curly top virus Eragrostis curvula streak virus Grapevine red blotch virus Maize streak virus Tomato pseudo-curly top virus Turnip curly top virus Citrus chlorotic dwarf associated virus Mulberry mosaic dwarf associated virus Commelina yellow mottle virus Cauliflower mosaic virus Cassava vein mosaic virus Petunia vein clearing virus Rose yellow vein virus Tobacco vein clearing virus Soybean chlorotic mottle virus Rice tungro bacilliform virus

Family Avsunviroidae

Genus Avsunviroid Elaviroid Pelamoviroid Apscaviroid Cocadviroid Coleviroid Hostuviroid Pospiviroid

Type species Avocado sunblotch viroid Eggplant latent viroid Peach latent mosaic viroid Apple scar skin viroid Coconut cadang-cadang viroid Coleus blumei viroid 1 Hop stunt viroid Potato spindle tuber viroid

(+) sense Unassigned ssDNA Viruses

Nanoviridae

(+/-) sense Unassigned ssDNA Viruses

Geminiviridae

Genus Unassigned Unassigned Unassigned Unassigned Unassigned Unassigned Unassigned Unassigned Unassigned Phytoreovirus Fijivirus Oryzavirus Babuvirus Nanovirus Unassigned Becurtovirus Begomovirus Capulavirus Curtovirus Eragrovirus Grablovirus Mastrevirus Topocuvirus Turncurtovirus Unassigned Unassigned

dsDNA (Reverse transcribing viruses)

Ortervirales

II. Plant viroids Nature of genome Order (+) sense ssRNA Unassigned Viroids Unassigned

Pospiviroidae

xxii

Introduction

The form and orthography of official virus nomenclature has thus changed over the years and continues to evolve. Tomato spotted wilt disease, which was first described in Australia in 1915, was identified as a virus in 1930 and named as tomato spotted wilt virus (Best 1968) and was first recognized as a species by the ICTV in the first report (1971) under the name Tomato spotted wilt virus of the Tomato spotted wilt virus group. This was amended in 1990 to the genus Tospovirus, family Bunyaviridae, and then the species name modified to Tomato spotted wilt tospovirus (still genus Tospovirus, family Bunyaviridae) in 2015. During 2016, and susequently in 2018b.v1 the species name changed to Tomato spotted wilt orthotospovirus, genus Orthotospovirus, family Tospoviridae, order Bunyavirales. The names of other viral species are also revised, with the genus name replacing the simple “virus” at the end of the species name, to immediately connect the species name with the appropriate genus. There are currently proposals to increase the number of taxonomic ranks to which viruses may be assigned, but there is a considerable range of opinion on this subject which may take years to resolve before a final decision is reached. Rather than describing the current and proposed rules of nomenclature and orthography here, the reader is therefore advised to refer for the current status and proposals, complete with current information on how to write a virus name, available at the ICTV website, https://talk.ictvonline.org/, which is the authoritative source of information on current virus taxonomy. Van Regenmortel (2001) has discussed the perspectives of binomial names for virus species.

Plant Viral Database It is necessary to have descriptions of the major parameters that identify a particular virus. As several viruses may infect a single plant species and the knowledge base of plant viruses is expanding constantly, efforts were made in the past to compile a global database of plant viruses. The Description of Plant Viruses (DPV) was originally published jointly by the Commonwealth Mycological Institute (CMI) and the Association of Applied Biologists (AAB), UK. The DPV series comprises of excellent descriptions of over 354 individual plant viruses that were originally published as hard copy between 1970 and 1989. Each DPV was prepared by an expert and contained cryptogram (identifying feature), synonyms, main disease symptoms, geographical distribution, host range, strains, transmission, serology, stability in sap, purification, properties of particles, particle structure and composition, relation with cells and tissues, notes, references, and a plate depicting diagnostic symptoms and virions. Subsequently, an online version of DPV was developed that since 1998 has included detailed descriptions of plant viruses together with information on taxonomy and sequences. It provides a comprehensive resource that is widely used for teaching, disease management, and research. The DPVweb (http://www.dpvweb.net) provides a source of information about viruses, viroids, and satellites of plants, fungi, and protozoa (Adams and Antoniw 2005, 2006). In the early 1980s, the Virus Ecology Research Group of the Research School of Biological Sciences, Australian National University, initiated work compiling a database on viruses of legumes to test the use of the computer-based Description Language for Taxonomy (DELTA) system (http://delta-intkey.com/ www/refs.htm) for storing and manipulating taxonomic descriptions (Boswell and Gibbs 1983). The project was subsequently extended to create a data bank for plant viruses, which was known as the Virus Identification Data Exchange (VIDE) project. The international VIDE project, which uses the DELTA database system, assembled diagnostic information about plant viruses from all over the world. This resulted in the first computer database of plant virus descriptions (Boswell et al. 1986). In 1991, it became the first component of the Universal Virus Database of the International Committee on Taxonomy of Viruses (ICTVdB) (Büchen-Osmond and Dallwitz 1996). The directory of the ICTVdB contains a list of approved virus names linked to virus descriptions coded from information in the seventh report of the ICTV (van Regenmortel et al. 2000b) and includes updates subsequently approved by the ICTV. It also

Introduction

xxiii

incorporates the plant virus database VIDEdB and is illustrated with EM pictures, diagrams, and images of symptoms contributed by virologists from around the world. The work of the ICTVdB, which was concluded in 2008, contained data on as many as 4949 virus species. Although the ICTV has discontinued the work on its database of virus descriptions, it has been suggested that the storage of metadata of viruses in public databases, such as Wikipedia (http://en.wikipedia.org) or the Encyclopedia of Life (http://www. eol.org) (Gibbs 2013), should be promoted. The VIDE database has been updated from time to time and is available online (http://pvo.bio-mirror.cn/refs.htm) (Brunt et al. 1996). The VIDE database became an important resource for plant viruses and was subsequently consolidated and published in the form of books by the CAB International: Viruses of Tropical Plants: Descriptions and Lists from the VIDE Database (Brunt et al. 1990) and Viruses of Plants: Descriptions and Lists from the VIDE Database (Brunt et al. 1996). The book, Viruses of Plants covered more than 900 viruses arranged alphabetically, with each described with characters related to the susceptibility of host plants and properties for virus identification. Detailed information on selected plant viruses, their images, maps, biology, distribution, impact, diagnosis, molecular biology, management, and bibliography are available online in the encyclopedia resource Invasive Species Compendium (http://www.cabi.org/isc) and Crop Protection Compendium (http://www.cabi.org/cpc) by CAB International. The European and Mediterranean Plant Protection Organization also provides datasheet information about plant viruses (https://www.eppo.int).

Books on Description of Plant Viruses The first book on plant viruses was published in 1937, A Textbook of Plant Virus Diseases, by K.M. Smith, which described plant viruses, their properties, and associated diseases. The science of plant virology grew rapidly due to the development of techniques to study protein and nucleic acids, the two major constituents of viruses. As a result, numerous plant viruses were discovered in various parts of the world. Over time several textbooks on plant viruses were written (Smith 1937; Gibbs and Harrison 1976; Matthews 1981; Sutic et al. 1999; Hull 2002; Loebenstein and Thottappilly 2003; Mahy and Van Regenmortel 2008; Rao et al. 2008, 2012; Subramanya Sastry 2013a, b; Subramanya Sastry and Zitter 2014; Mandal et al. 2017). Books on individual virus description were written during the 1990s: Viruses of Tropical Plants: Descriptions and Lists from the VIDE Database and Viruses of Plants (Brunt et al. 1990, 1996). The generation of plant virus genome sequence information provided the single most important criterion for the identification and differentiation of plant viruses. As a result, the confusion over the synonymous identity of several plant viruses was resolved, and in the ninth report of the ICTV, definite plant virus species as well as tentative members of a genus were listed (King et al. 2012). However, a basic description of the officially recognized plant viruses is not available.

The Design and Objectives of the Book The available books that describe plant viruses were published 20 years ago. During the last two decades, numerous plant viruses and viroids have been discovered, and their nomenclature and classifications have changed over time. In this Encyclopedia of Plant Viruses and Viroids, the natural infection of different viruses and viroids on nearly 1020 host plants are described based on the current international taxonomic criteria of the ICTV (Anon 2017). The book has been designed with the objective of providing a comprehensive description of plant viruses and viroids known to infect a specific plant species globally; entries are arranged by the alphabetical order of plant species, and all the viruses and viroids characterized to date have been included under each plant species. Most of the previously published books described viruses

xxiv

Introduction

in alphabetical order, which did not provide a ready reference of the number of viruses or viroids known to infect a particular plant species. All of the virus species that have been recognised by ICTV, have been included; in addition tentative species, which have recently been published or published previously but not yet recognised by ICTV for various reasons, have been included in this encyclopaedia. The virus name in this book appears as the major subtitle in each section of virus description. In order to differentiate the officially recognised and tentative virus species, they were presented as italic and non-italic subtitle. As the subtitle is a name of a virus which is necessary to refer to the subsequent text, it has been abbreviated for convenience, even though it is an official name (virus species). Each virus has been described with the following descriptors (to the extent that information is available): taxonomic position, geographical distribution, symptoms and host range, transmission, virion properties, and genome properties. A consolidated list of references cited in the text has been provided at the end of the virus descriptions under each host.

References Adams MJ, Antoniw JF (2005) DPVweb: an open access internet resource on plant viruses and virus diseases. Outlook Pest Manag 16:268–270 Adams MJ, Antoniw JF (2006) DPVweb: a comprehensive database of plant and fungal virus genes and genomes. Nucleic Acids Res 34(Database issue):D382–D385 Adams MJ, Lefkowitz EJ, King AM, Harrach B, Harrison RL, Knowles NJ, Kropinski AM, Krupovic M, Kuhn JH, Mushegian AR, Nibert ML, Sabanadzovic S, Sanfaçon H, Siddell SG, Simmonds P, Varsani A, Zerbini FM, Orton RJ, Smith DB, Gorbalenya AE, Davison AJ (2017) 50 Years of the International Committee on Taxonomy of Viruses: progress and prospects. Arch Virol 162:1441–1446 Anon (2017) Virus taxonomy: the classification and nomenclature of viruses. The online (10th) report of the ICTV. https://talk.ictvonline.org/ictv-reports/ictv_online_report/ Best RJ (1968) Tomato spotted wilt virus. In: KM Smith and MA Lauffer (eds). Advances in Virus Research, Vol. 13. Academic Press, New York. pp 66–146 Boswell KF, Gibbs AJ (eds) (1983) Viruses of legumes 1983 – Descriptions and keys from VIDE. Australian National University, Canberra, 139 pp Boswell KF, Dallwitz MJ, Gibbs AJ, Watson L (1986) The VIDE (Virus Identification Data Exchange) project: a data bank for plant viruses. Rev Plant Pathol 65:221–231 Brandes J, Wetter C (1959) Classification of elongated plant viruses on the basis of particle morphology. Virology 8:99–115 Brunt A, Crabtree K, Gibbs A (1990) Viruses of tropical plants: Descriptions and lists from the VIDE database. CAB International, Wallingford, 707 pp Brunt AA, Crabtree K, Dallwitz MJ, Gibbs AJ, Watson L (1996) Viruses of plants. Descriptions and lists from the VIDE database. CAB International, Wallingford, 1484 pp Büchen-Osmond C, Dallwitz M (1996) Towards a universal virus database – progress in the ICTVdB. Arch Virol 141:392–399 Calisher CH, Horzinek M, Mayo MA, Ackermann HW, Maniloff (1995) Sequence analyses and unifying system of virus taxonomy: consensus via consent. Arch Virol 140:2093–2099 Diener TO (1971) Potato spindle tuber ‘virus’. IV. A replicating, low molecular weight RNA. Virology 45:411–428 Fauquet C, Mayo MA, Maniloff J, Desselberger U, Ball LA (eds) (2005) Virus taxonomy: eighth report of the International Committee on Taxonomy of Viruses. Elsevier/Academic, Amsterdam

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Fenner F (1976) Classification and nomenclature of viruses. Second report of the International Committee on Taxonomy of Viruses. Intervirol 7:1–115 Flores R, Di Serio F, Hernandez C (1997) Viroids: the non-encoding genomes. Seminars Virol 8:65–73 Francki RIB, Fauquet CM, Knudson DL, Brown F (1991) Classification and nomenclature of viruses. Fifth report of the International Committee on Taxonomy of Viruses. Arch Virol Suppl. 2. Springer, Vienna Gibbs AJ (1968) Cryptograms. In: Martyn EB (ed). Plant Virus Names. Commonwealth Mycol Inst Kew, Surrey, pp 135–149 Gibbs AJ (2013) Viral taxonomy needs a spring clean; its exploration era is over. Virology J 10:254 Gibbs AJ, Harrison BD (1968) Realistic approach to virus classification and nomenclature. Nature 218: 927–929 Gibbs A, Harrison BD (1976) Plant Virology: The Principles. Edward Arnold, London (UK). ISBN: 071312508X, 292 pp. Gibbs AJ, Harrison BD, Watson DH, Wildy P (1966) What's in a virus name? Nature 209:450–454 Holmes FO (1939) Proposal for extension of the binomial system of nomenclature to include viruses. Phytopathology 29:431–436 Hull R (2002) Matthews’ Plant Virology, 4th edn, New York, NY: Academic Press. 1056 p, ISBN: 9780-12-361160-4 Johnson J (1927) The classification of plant viruses. Res Bull Wis Agric Exp Stn 76:1–16 Johnson J, Hoggan IA (1935) A descriptive key for plant viruses. Phytopathology 25:328–343 King AMQ, Adams MJ, Carstens EB, Lefkowitz EJ (2012) Virus taxonomy: classification and nomenclature of viruses. Ninth report of the International Committee on Taxonomy of Viruses. Elsevier Academic Press, San Diego Loebenstein G, Thottappilly G (2003) Virus and virus-like diseases of major crops in developing countries. Springer Publishers, Netherlands. https://doi.org/10.1007/978-94-007-0791-7 Mahner M, Bunge M (1997) Foundations of biophilosophy. Springer, Berlin Mahy BWJ, Van Regenmortel MHV (2008) Desk encyclopaedia of plant and fungal virology. Elsevier Publishers Mandal B, Rao GP, Baranwal VK, Jain RK (2017) A century of plant virology in India. Springer Publishers, p 805 Matthews RE (1979) Third report of the International Committee on Taxonomy of Viruses. Classification and nomenclature of viruses. Intervirol 12:129–296 Matthews REF (1981) Plant Virology, 2nd edn. Academic Press, ISBN: 9781483258669, pp 916 Matthews REF (1982) Classification and nomenclature of viruses. Fourth report of the international committee on taxonomy of viruses. Intervirol 17:1–199 Murphy FA, Fauquet CM, Bishop DHL, Ghabrial SA, Jarvis AW, Martelli GP, Mayo MA, Summers MD (1995) Virus taxonomy. Sixth report of the International Committee on Taxonomy of Viruses. Springer, Vienna Pringle CR (1991) The 20th meeting of the executive committee of the ICTV. Virus species, higher taxa, a universal database and other matters. Arch Virol 119:303e4.5 Rao GP, Kumar PL, Holguin-Pena RJ (2008) Characterization, diagnosis and management of plant viruses. Vegetable and pulse crops, vol III. Studium Press, LLS, Houstan, 408 p Rao GP, Baranwal VK, Mandal B, Rishi N (2012) Recent trends in plant virology. Studium Press LLC, USA, 493 p Smith KM (1937) Text book of Plant Virus Diseases. P Blakiston’s Son and Company, Philadelphia Simmonds P (2015) Methods for virus classification and the challenge of incorporating metagenomic sequence data. J Gen Virol 96:1193–1206

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Stobbe AH, Daniels J, Espindola AS, Verma R, Melcher U, Ochoa-Corona F et al (2013) E-probe diagnostic nucleic acid analysis (EDNA): a theoretical approach for handling of next generation sequencing data for diagnostics. J Mirobiol Methods 94:356–366 Subramanya Sastry K (2013a) Seed-borne plant virus diseases. Springer Publishers, p 327 Subramanya Sastry K (2013b) Plant virus and viroid diseases in the tropics. Introduction of plant viruses and sub-viral agents, classification, assessment of loss, transmission and diagnosis, vol 1. Springer Publishers, 361 p Subramanya Sastry K, Zitter TA (2014) Plant virus and viroid diseases in the tropics. Epidemiology and management of plant viruses, vol 2. Springer Publishers, 489 p Sutic DE, Ford RE, Tosic MT (1999) Handbook of plant virus diseases. CRC Press, pp 584 Tsugita A, Gish DT, Young J, Fraenkel-Conrat H, Knight CA, Stanley WM (1960) The complete amino acid sequence of the protein of tobacco mosaic virus. Proc Natl Acad Sci U S A 46:1463–1469 Van Regenmortel MH (1989) Applying the species concept to plant viruses. Arch Virol 104:1–17 Van Regenmortel MHV (1990) Virus species, a much overlooked but essential concept in virus classification. Intervirol 31:241–254 Van Regenmortel MHV (2001) Perspectives on binomial names for virus species. Arch Virol 146:1637–1640 Van Regenmortel MHV (2016a) Only viruses, but not their genome sequences, can be classified into hierarchical species and genus classes. Curr Top Virol 13:59–68 Van Regenmortel MHV (2016b) “Classes, taxa and categories in hierarchical virusclassification: a review of current debates on definitions and names of virus species” appeared in Bionomina. Dumerilia 10:1e20 Van Regenmortel MHV (2016c) Recent developments in the definition and official names of virus species. In: Genetics and Evolution of Infectious Diseases, pp 1–23. https://doi.org/10.1016/B9780-12-799942-5.00001-9 Van Regenmortel MHV, Fauquet CM (2002) Only italicized species names of viruses have a taxonomic meaning. Arch Virol 147:2247–2250 Van Regenmortel MHV, Bishop DHL, Fauquet CM, Mayo MA, Maniloff J, Calisher CH (1997) Guidelines to the demarcation of virus species. Arch Virol 142:1505–1518 Van Regenmortel MHV, Mayo MA, Fauquet CM, Maniloff J (2000a) Virus nomenclature: consensus versus chaos. Arch Virol 145:2227–2232 Van Regenmortel MHV, Fauquet CM, Bishop DHL, Carstens EB, Estes MK, Lemon SM, Maniloff J, Mayo MA, McGeoch DJ, Pringle CR, Wickner RB (2000b) Virus taxonomy: Seventh report of the International Committee on Taxonomy of Viruses. Academic, San Diego Wildy P (1971) Classification and nomenclature of viruses. First report of the International Committee on Nomenclature of Viruses. Monog Virol 5:1–81

About the Authors

Dr. K. Subramanya Sastry a retired Principal Scientist from Indian Council of Agricultural Research (ICAR), India, has carried out research in plant virology for over 28 years. During his service period, he has served as scientist at different ICAR institutions, such as IIHR, Bengaluru; IIOR (DOR) Hyderabad; and IIMR (NRCS), Hyderabad (India). Prof. Sastry’s research has been primarily on epidemiology, management, and molecular and biotechnological approaches for characterization of viruses and virus-like diseases of crops of horticultural, oil seeds, and millets. He has published 83 research articles and 5 reference books on plant virology, and 3 of them are through Springer Publishers. Dr. K. Subramanya Sastry is the editor-inchief of this present reference book entitled Encyclopedia of Plant Viruses and Viroids.

Dr. Bikash Mandal is currently working as Principal Scientist in Advanced Center for Plant Virology, Indian Agricultural Research Institute (IARI), New Delhi. His research and teaching area is plant xxvii

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About the Authors

virology. He has totally 153 publications that include 2 books, 8 book chapters, 72 research papers, 5 popular technical articles, and 65 conference abstracts. He is the editor-in-chief of the journal VirusDisease (Springer) and Virus Research News (Newsletter).

Dr. John Hammond is currently working as Research Plant Pathologist, United States Department of Agriculture, Agricultural Research Service, Floral and Nursery Plants Research Unit, Beltsville. His major areas of research are viruses affecting ornamental crops, with emphasis on potyvirus, potexvirus, and carlavirus detection, differentiation, and methods of introducing resistance, use of transgenic plants to examine virus resistance and infectious viral clones to determine factors affecting host range, symptom induction, and systemic movement, and development of microarrays for plant virus detection and identification.

Dr. Simon W. Scott has recently retired from Clemson University as a Professor Emeritus in Plant Pathology. His major areas of research were viruses and virus-like agents affecting woody deciduous species with an emphasis on viruses that affect the dominant fruit crop in South Carolina (peaches). He has established a program to index large blocks of peach trees in the southeastern USA prior to propagation as part of the USDA/APHIS National Clean Plant Network. In addition, he has produced extensive sequence data for a number of Ilarviruses, allowing long-standing taxonomic anomalies to be corrected.

About the Authors

xxix

Dr. Robert William Briddon has worked as Principle Investigator in Agricultural Biotechnology Division, National Institute of Biotechnology and Genetic Engineering (funded by United States Department of Agriculture (USDA) through the International Center for Agricultural Research in the Dry Areas (ICARDA, Islamabad Office). His research area is in study of vector transmission of plant infecting viruses, particularly the interactions involved between virus and insect vector of circulatively and propagatively transmitted viruses and the evolution thereof. He has totally 135 publications in ISI Web of knowledge. He is also a member of British Society for Plant Pathology, European Whitefly Study Network, and International Committee on Taxonomy of Viruses.

A

Abelia spp. Family: Caprifoliaceae

Eggplant mosaic virus

Trees/Shrubs

(EMV)

Synonyms Abelia latent virus Taxonomic position Genus: Tymovirus

Family: Tymoviridae

EMV isolated from symptomless Abelia grandiflora in the USA was initially identified as Abelia latent virus (Waterworth et al. 1975) but is now recognized as a strain of Eggplant mosaic virus. The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of EMV, refer to Solanum melongena.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Abelia x grandiflora was reported from Georgia, USA (Ruter and Gitaitis 1993). The virus-infected abelia plants do not exhibit any external symptoms. The virus is transmitted by thrips vectors in a persistent, propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

References Ruter JM, Gitaitis RD (1993) Impatiens necrotic spot virus in woody landscape plants in Georgia. Plant Dis 77:318 Waterworth HE, Kaper JM, Koenig R (1975) Purification and properties of a Tymovirus from Abelia. Phytopathology 65:891–896 © Springer Nature India Private Limited 2019 K. S. Sastry et al., Encyclopedia of Plant Viruses and Viroids, https://doi.org/10.1007/978-81-322-3912-3

2

Abelmoschus esculentus (Bhendi, Ladies’ finger, Okra)

Abelmoschus esculentus (Bhendi, Ladies’ finger, Okra) Synonyms Hibiscus esculentus Family: Malvaceae

Vegetable

Bhendi yellow vein Bhubhaneswar virus Taxonomic position Genus: Begomovirus

(BYVBhV)

Family: Geminiviridae

Geographical distribution BYVBhV infection in plants of Hibiscus esculentus was reported from Bhubhaneswar (India) (Venkataravanappa et al. 2013a). Symptoms and host(s) BYVBhV-infected okra plants exhibit prominent yellow vein mosaic symptoms and stunting. The virus was isolated from okra (Abelmoschus esculentus) but no further investigation of the host range of the virus has been conducted. Transmission The virus has been shown to be transmitted by the whitefly vector of begomoviruses, Bemisia tabaci (Venkataravanappa et al. 2013a). Although the mechanism of transmission of BYVBhV by B. tabaci has not been investigated, the mechanism is likely to be circulative, non-propagative in common with all begomoviruses. Virion properties and genome The structure of BYVBhV particles has not been investigated. In common with all geminiviruses, the virions of BYVBhV are likely geminate (twinned icosahedra). Only a single isolate of BYVBhV has been identified and characterized to date. The evidence suggests that this virus has a monopartite genome. The genome sequence comprises 2757 nt (FJ589571 = NC_012041) (Venkataravanappa et al. 2013a). The characterized genome of BYVBhV encodes the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated. BYVBhV has been shown to associate with a betasatellite (Briddon 2001; Venkataravanappa et al. 2013a; Brown et al. 2015; Zerbini et al. 2017).

Bhendi yellow vein Haryana virus Taxonomic position Genus: Begomovirus

(BYVHaV)

Family: Geminiviridae

Abelmoschus esculentus (Bhendi, Ladies’ finger, Okra)

3

Geographical distribution BYVHaV infection in plants of Abelmoschus esculentus was reported from Haryana, India (Venkataravanappa et al. unpublished - FJ561298). Symptoms and host(s) The virus-infected okra plants exhibit yellow vein symptoms. Transmission The virus transmission has not been reported but is expected to be transmitted by the whitefly, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2740 nt (FJ561298) (Briddon 2001; Brown et al. 2015).

Bhendi yellow vein mosaic Delhi virus Taxonomic position Genus: Begomovirus

(BYVMDV)

Family: Geminiviridae

Geographical distribution BYVMDV infection in plants of Abelmoschus esculentus was reported from New Delhi, India (Venkataravanappa et al. 2012a). Symptoms and host(s) The virus-infected bhendi plants show yellow vein mosaic, vein twisting, reduced leaves, and a bushy appearance. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2751 nt (FJ515747 = NC_011919) (Venkataravanappa et al. 2012a; Brown et al. 2015; Zerbini et al. 2017).

Bhendi yellow vein mosaic virus Taxonomic position Genus: Begomovirus

(BYVMV)

Family: Geminiviridae

Geographical distribution BYVMV disease was first reported in Abelmoschus esculentus from Mumbai in India (Kulkarni 1924).

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Abelmoschus esculentus (Bhendi, Ladies’ finger, Okra)

BYVMV has been identified in India, Sri Lanka, Pakistan, Chad, Cote D’Ivoire, Nigeria, China, Sudan, and Thailand (Pun and Doraiswamy 1999; Mansoor et al. 2001; Jose and Usha 2003; Usha 2008; Rishi 2009; Xie et al. 2010; Tsai et al. 2013; Sayed et al. 2014; Kumar et al. 2017; Tharmila et al. 2017). The virus has also been known under the names okra yellow vein mosaic virus, bhendi yellow vein mosaic Maharashtra virus, bhendi yellow vein India virus, okra leaf curl virus, and bhendi yellow vein Delhi virus (Zhou et al. 1998).

Symptoms and host(s) The virus-infected okra plants exhibit yellowing of leaf veins. The yellow network of veins is very conspicuous, and veins and veinlets are thickened. In severe cases, the chlorosis may extend to the interveinal area and may result in complete yellowing of leaves. Fruits are dwarfed, malformed, and yellow green.

Transmission The virus has been shown to be transmitted by the whitefly Bemisia tabaci (Varma 1952; Jose and Usha 2003; Venkataravanappa et al. 2012a). Although, the mechanism of transmission of BYVMV by B. tabaci has not been investigated, the mechanism is likely to be circulative, non-propagative in common with all begomoviruses. Experimentally, the virus has been transmitted to okra by Agrobacterium-mediated inoculation of cloned virus in the presence of a betasatellite (Jose and Usha 2003).

Virion properties and genome The structure of BYVMV particles has not been investigated. In common with all geminiviruses, the virion of BYVMV has a geminate morphology (twinned quasi-icosahedra). For the vast majority of isolates of BYVMV that have been characterized, the genome consists of a single circular, single-stranded DNA of 2741 nt (AF241479 = NC_003418). These monopartite isolates are generally associated with a betasatellite, which is required for infection of okra (Jose and Usha 2003). Several isolates of BYVMV have been identified which appear to be bipartite, consisting of two single circular, single-stranded DNA components. However, Koch’s postulates have been satisfied only for BYVMV causing yellow vein mosaic disease in okra in the presence of a betasatellite (Jose and Usha 2003). This is not the case for the bipartite BYVMV (Venkataravanappa et al. 2012a, 2015b). It is important to note that the DNA B component associated with the apparently bipartite BYVMV isolates showed high levels of sequence identity to the DNA B components of Tomato leaf curl New Delhi virus, a bipartite virus with a wide host range, which also has been reported in okra (Venkataravanappa et al. 2012a, 2014). The database accession numbers for the genome of DNA A are AJ002453, FJ515747, JQ326263, and GU112049 and DNA B are HQ542082, HQ586005, and JQ326263 (Briddon 2001; Jose and Usha 2003; Ghosh et al. 2008; Zhou 2013; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes/DNA A component of BYVMV encodes the six genes that are typically encoded by monopartite begomoviruses and the DNA A component of bipartite begomoviruses originating from the Old World, two in the virion-sense and four in the complementary-sense. The expression and function of these genes have not been investigated for BYVMV. The monopartite form of the virus has been shown to associate with a betasatellite (Jose and Usha 2003).

Abelmoschus esculentus (Bhendi, Ladies’ finger, Okra)

Chino del tomate virus

(CdTV)

Synonyms Okra yellow mottle Iguala virus Taxonomic position Genus: Begomovirus

5

(OYMoIgV)

Family: Geminiviridae

OYMoIgV infection in plants of Hibiscus esculentus was reported from Mexico (Valadez-Moctezuma et al. unpublished - AY751753). The virus-infected okra plants exhibit yellow mottle symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of CdTV, refer to Solanum lycopersicum.

Chickpea chlorotic dwarf virus Taxonomic position Genus: Mastrevirus

(CpCDV)

Family: Geminiviridae

CpCDV infection in plants of Abelmoschus esculentus was reported from Pakistan (Zia-Ur-Rehman et al. 2017). The virus-infected plants exhibit leaf curling, yellowing, and vein thickening symptoms. The virus is transmitted by leafhopper vectors in a persistent, circulative, and non-propagative manner, and infects a number of economically important crops. The virus is not transmitted by mechanical inoculation. For more details of CpCDV, refer to Cicer arietinum.

Cotton leaf curl Alabad virus Taxonomic position Genus: Begomovirus

(CLCuAlV)

Family: Geminiviridae

CLCuAlV infection in plants of Abelmoschus esculentus was reported from North India (Venkataravanappa et al. 2012b). The virus-infected okra plants exhibit yellow vein mosaic, downward curling and vein twisting symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-persistent manner. The virus is not transmissible by mechanical sap-inoculation, and also not by contact between plants. For more details of CLCuAlV, refer to Gossypium spp.

Cotton leaf curl Bangalore virus Taxonomic position Genus: Begomovirus

(CLCuBaV)

Family: Geminiviridae

CLCuBaV infection in plants of Abelmoschus esculentus was reported from India (Venkataravanappa et al. 2013b). The virus-infected okra plants exhibit upward leaf curling, vein-clearing, vein-thickening, and yellowing symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-persistent manner. The virus is not transmissible by mechanical sap-inoculation, and also not by contact between plants. For more details of CLCuBaV, refer to Gossypium spp.

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Cotton leaf curl Gezira virus

(CLCuGeV)

Synonyms Okra leaf curl Cameroon virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

CLCuGeV is one of the serious viral pathogens in Abelmoschus esculentus in Burkina Faso, Pakistan, Cameroon, Egypt, Oman, Mali, Sudan, Cote d’Ivoire, and Niger (Kon et al. 2009; Shih et al. 2009; Tiendrebeogo et al. 2010; Leke et al. 2013; Sattar 2012; Akhtar et al. 2014; Idris et al. 2014). The virus causes leaf curl disease in okra. The veins of the leaves are dark green and swollen, and severely affected plants are highly stunted and unproductive. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by mechanical sap-inoculation, and also not by contact between plants. For more details of CLCuGeV, refer to Gossypium spp.

Okra enation leaf curl virus Taxonomic position Genus: Begomovirus

(OELCuV)

Family: Geminiviridae

Geographical distribution OELCuV infection in plants of Abelmoschus esculentus was reported from India and Pakistan (Singh and Dutta 1986; Chakraborty et al. 1997; Chandran et al. 2013; Venkataravanappa et al. 2012a; Serfraz et al. 2015; Shruti et al. 2016). Symptoms and host(s) The virus-infected okra plants exhibit leaf curling, crumpling, and enations on the underside of the leaves. The virus has been isolated from okra, papaya, and cotton. Transmission In common with all viruses of the genus Begomovirus, OELCuV is transmitted by the whitefly Bemisia tabaci (Venkataravanappa et al. 2015a), although the mechanism of transmission of OELCuV by the whitefly has not been investigated. Experimentally, OELCuV has been whitefly transmitted to Abelmoschus esculentus (okra), Althaea rosea (hollyhock), Datura stramonium, Nicotiana clevelandii, N. glutinosa, N. occidentalis, N. sylvestris, N. tabacum, and N. benthamiana. In each case the virus induced leaf curling and foliar enations (Venkataravanappa et al. 2015a). The virus is not transmitted by mechanical sap-inoculation, and not through seed or aphids. Virion properties and genome The structure of the virions of OELCuV has not been investigated. In common with all geminiviruses, the virions are presumed to have geminate morphology (twinned quasi-icosahedra). OELCuV is a typical monopartite begomovirus. The virus genome consists of a single circular molecule of single-stranded DNA of ~2724 nt (GU111998 = NC_014894). The genomic component of OELCuV encodes the six genes typically encoded by monopartite begomoviruses originating from the

Abelmoschus esculentus (Bhendi, Ladies’ finger, Okra)

7

Old World, two in the virion-sense and four in the complementary-sense. The expression and function of the genes have not been investigated so far. Most isolates of OELCuV have been shown to associate with a betasatellite (Briddon 2001; Venkataravanappa et al. 2011, 2012a, 2015a; Brown et al. 2015; Zerbini et al. 2017).

Okra leaf curl Oman virus Taxonomic position Genus: Begomovirus

(OLCOV)

Family: Geminiviridae

Geographical distribution OLCOV infection in plants of Abelmoschus esculentus was reported from Oman (Akhtar et al. 2014). Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm, with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2788 nt (HE862273 = NC_028985) (Briddon 2001; Akhtar et al. 2014; Brown et al. 2015; Zerbini et al. 2017).

Okra mosaic virus Taxonomic position Genus: Tymovirus

(OkMV)

Family: Tymoviridae

Geographical distribution OkMV infection in plants of Abelmoschus esculentus was initially reported in Cote d’Ivoire (Givord et al. 1972). The virus spreads in West Africa and Brazil (Igwegbe 1983; Atiri 1984; Fauquet and Thouvenel 1987; Fajinmi and Fajinmi 2010). Symptoms and host(s) Typical symptoms of OkMVon okra include vein chlorosis, vein-banding, mosaic, and stunting. Yields of affected crops are greatly reduced. The natural hosts of OkMV include jute mallow (Corchorus olitorius), roselle (Hibiscus sabdariffa), hibiscus (H. rosa-sinensis), and West African okra (Abelmoschus caillei). Transmission The virus is transmitted by a beetle vector, Podagrica decolorata in Cote d’Ivoire, Podagrica uniforma, and P. sjostedti in a semi-persistent manner in Nigeria (Givord and den Boer 1980; Lana and AjibolaTaylor 1976). A report of whitefly, Bemisia tabaci, as a vector in Nigeria needs confirmation. The virus is not transmitted by Aphis gossypii, Chrysolagria cuprina, Lagria villosa, Medythia quaterna, Ootheca mutabilis, and Nisotra dilecta. The virus is mechanically sap-transmissible and also by grafting, but not transmitted by seed.

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Abelmoschus esculentus (Bhendi, Ladies’ finger, Okra)

Virion properties and genome The virions are non-enveloped, isometric capsid with T=3 icosahedral symmetry, about 30 nm in diameter. The genome consists of a single molecule of positive sense, ssRNA of 6223 nt (EF554577 = NC_009532). The 30 -terminus has a tRNA-like structure (Givord and Hirth 1973; Martelli et al. 2002; Stephan et al. 2008).

Okra mottle virus Taxonomic position Genus: Begomovirus

(OMoV)

Family: Geminiviridae

Geographical distribution OMoV infection in plants of Hibiscus esculentus was reported from Brazil (Albuquerque et al. 2013). Symptoms and host(s) The virus-infected okra plants exhibit mottling and chlorotic spot symptoms. The virus has also been isolated from soybean (Glycine max), although the symptoms associated with infected plants were not detailed. The host range of the virus has not been investigated. Transmission The transmission of OMoV has not been investigated. It is likely that, in common with other begomoviruses, OMoV will be transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Virion properties and genome The structure of the virions of OMoV has not been investigated. In common with all geminiviruses, the virions of OMoV are likely geminate (twinned icosahedra). OMoV is typical of bipartite begomoviruses native to the New World. The genome consists of two circular, single-stranded DNA components, DNA A and DNA B of 2660 nt (EU914817 = NC_011181; FJ686695) and 2653 nt (EU914818 = NC_011182; FJ686696) respectively (Albuquerque et al. 2013). Typical of the majority of bipartite begomoviruses native to the New World, the DNA A component of OMoV encodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA B encodes one gene in each orientation. The expression and function of these genes have not been investigated for OMoV (Briddon 2001; Albuquerque et al. 2013; Brown et al. 2015; Zerbini et al. 2017).

Okra yellow crinkle virus

(OYCrV)

Synonyms Okra yellow crinkle Mali virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Abelmoschus esculentus (Bhendi, Ladies’ finger, Okra)

9

Geographical distribution OYCrV infection in plants of Abelmoschus esculentus was first recorded in Mali (Shih et al. 2007) and thereafter reported from Cameroon and Cote d’Ivoire (Leke et al. 2013; Seka et al. 2016). Symptoms and host(s) The virus-infected okra plants exhibit foliar yellowing, crinkling, and cupping symptoms. Transmission The transmission of OYCrV has not been investigated. It is likely that, in common with other begomoviruses, OYCrV is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of OYCrV has not been investigated. In common with all geminiviruses, the virions of OYCrV are likely geminate (twinned quasi-icosahedra). OYCrV is a typical Old World monopartite begomovirus. The genome of OYCrV consists of a single circular molecule of single-stranded DNA of ~2792 nt (DQ875879 = NC_008377). The expression and function of the genes have not been investigated for OYCrV. OYCrV associates with betasatellites (Briddon 2001; Shih et al. 2007; Leke et al. 2013; Brown et al. 2015; Zerbini et al. 2017).

Okra yellow mosaic Mexico virus Taxonomic position Genus: Begomovirus

(OYMMV)

Family: Geminiviridae

Geographical distribution OYMMV infection in plants of Abelmoschus esculentus was reported from the USA and Mexico (De La Torre-Almaraz et al. 2006; Hernandez-Zepeda et al. 2010). Symptoms and host(s) The virus-infected okra plants exhibit symptoms of irregular yellow patches on leaves and distinctive yellow borders on leaf edges and chlorosis of subsequently developing leaves. The virus has also been isolated from a weed (unidentified Sida spp.), although the symptoms were not specified. The host range of the virus has not been investigated. Transmission The transmission of OYMMV has not been investigated. It is likely that, in common with other begomoviruses, the virus is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of OYMMV has not been investigated. In common with all geminiviruses, the virions of OYMMV are likely geminate (twinned icosahedra). OYMMV is typical of bipartite begomoviruses native to the New World. The genome consists of two circular, single-stranded DNA components, DNA A of 2612 nt (DQ022611 = NC_014066; GU990612, GU990613, GU990614, HM035059, HQ020409, HQ116414) and DNA B of 2,594 nt (GU972604, HM035060) (Hernandez-Zepeda et al. 2010). Typical of the majority of bipartite

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begomoviruses native to the New World, the DNA A component of OYMMV encodes five genes, two in the virion-sense and four in the complementary-sense, whereas the DNA B encodes one gene in each orientation. The expression and function of these genes have not been investigated for OYMMV (Briddon 2001; Hernandez-Zepeda et al. 2010; Brown et al. 2015; Zerbini et al. 2017).

Radish leaf curl virus Taxonomic position Genus: Begomovirus

(RaLCuV)

Family: Geminiviridae

RaLCuV infection in plants of Abelmoschus esculentus was reported from India (Kumar et al. 2012). The virus-infected okra plants exhibit leaf curling and overall stunting of plants that bore no fruit. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of RaLCuV, refer to Raphanus sativus.

Sida micrantha mosaic virus Taxonomic position Genus: Begomovirus

(SiMMV)

Family: Geminiviridae

SiMMV infection in plants of Abelmoschus esculentus was reported from Brazil (Aranha et al. 2011; Inoue-Nagata et al. 2016). The virus-infected okra plants exhibit mosaic and chlorotic spot symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of SiMMV, refer to Sida spp.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Abelmoschus esculentus was reported from India (Krishnareddy et al. 2003; Rajamanickam and Karthikeyan 2014). The typical symptoms on okra displaying fruit distortion disease due TSV are chlorotic spots, chlorotic leaf blotches, distortion of leaves, chlorotic streaking, and distortion of fruits. Orange discoloration of terminal young leaves has also been noticed. The virus is transmitted by thrips vectors, the virus present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible. The virus is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

Abelmoschus esculentus (Bhendi, Ladies’ finger, Okra)

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TuMV infection in plants of Hibiscus esculentus was reported from Israel (Gera et al. 2001). The virusinfected okra plants exhibit symptoms of chlorosis, vein-clearing, necrosis, and growth reduction. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a number of hosts. For more details of TuMV, refer to Brassica rapa.

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Kumar J, Kumar A, Singh SP, Roy JK, Lalit A, Parmar D, Sharma NC, Tuli R (2012) First report of Radish leaf curl virus infecting okra in India. New Dis Rep 25:9 Kumar RV, Prasanna HC, Singh AK, Ragunathan D, Garg GK, Chakraborty S (2017) Molecular genetic analysis and evolution of begomoviruses and betasatellites causing yellow mosaic disease of bhendi. Virus Genes 53:275–285 Lana AO, Ajibola-Taylor T (1976) The insect transmission of an isolate of Okra mosaic virus occurring in Nigeria. Ann Appl Biol 82:361–364 Leke WN, Sattar MN, Ngane EB, Ngeve JM, Kvarnheden A, Brown JK (2013) Molecular characterization of begomoviruses and DNA satellites associated with Okra leaf curl disease in Cameroon. Virus Res 174:116–125 Mansoor S, Amin I, Hussain M, Zafar Y, Bull S, Briddon RW, Markham PG (2001) Association of a disease complex involving a Begomovirus, DNA 1 and a distinct DNA ß with leaf curl disease of okra in Pakistan. 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In: International conference (VIROCON 2016), Bangalore, p 143 Singh SJ, Dutta OP (1986) Enation leaf curl of okra – a new virus disease. Indian J Virol 2:114–117 Stephan D, Siddiqua M, Hoang AT, Engelmann J, Winter S, Maiss E (2008) Complete nucleotide sequence and experimental host range of Okra mosaic virus. Virus Genes 36:231–240 Tharmila CJ, Jeyaseelan EC, Ihsan U, Wetten AC, De Costa DM, Shaw MW (2017) First report on association of Okra yellow vein mosaic virus with yellow vein mosaic disease of okra (Abelmoschus esculentus) in Sri Lanka. Plant Dis 101:1335 Tiendrebeogo F, Lefeuvre P, Hoareau M, Villemot J, Konaté G, Traoré AS, Barro N, Traoré VS, Reynaud B, Traoré O, Lett JM (2010) Molecular diversity of Cotton leaf curl Gezira virus isolates and their satellite DNAs associated with okra leaf curl disease in Burkina Faso. Virol J 7:48 Tsai WS, Shih SL, Lee LM, Wang JT, Duangsong U, Kenyon L (2013) First report of Bhendi yellow vein mosaic virus associated with yellow vein mosaic of okra (Abelmoschus esculentus) in Thailand. Plant Dis 97:291 Usha R (2008) Bhendi yellow vein mosaic virus. In: Rao GP, Kumar PL, Holguin-Pena RJ (eds) Characterization, diagnosis and management of plant viruses, vol 3. Studium Press, Houston, pp 387–392 Varma PM (1952) Studies on the relationship of the Bhendi yellow vein mosaic virus and its vector, the whitefly (Bemisia tabaci Gen.). Indian J Agric Sci 22:75–91 Venkataravanappa V, Lakshminarayana Reddy CN, Swaranalatha P, Jalali S, Briddon R, Krishna Reddy M (2011) Diversity and phylogeography of begomovirus-associated beta satellites of okra in India. Virol J 8:555 Venkataravanappa V, Lakshminarayana Reddy CN, Jalali S, Krishna Reddy M (2012a) Molecular characterization of distinct bipartite Begomovirus infecting bhendi (Abelmoschus esculentus L.) in India. Virus Genes 44(3):522–535 Venkataravanappa V, Lakshminarayana Reddy CN, Swarnalatha P, Devaraju JS, Krishna Reddy M (2012b) Molecular evidence for association of Cotton leaf curl Alabad virus with yellow vein mosaic disease of okra in North India. Arch Phytopathol Plant Protect 45:2095–2113 Venkataravanappa V, Lakshminarayana Reddy CN, Jalali S, Krishna Reddy M (2013a) Molecular characterization of a new species of Begomovirus associated with yellow vein mosaic of bhendi (Okra) in Bhubhaneswar, India. Eur J Plant Pathol 136:811–822 Venkataravanappa V, Lakshminarayana Reddy CN, Devaraju A, Jalali S, Krishna Reddy M (2013b) Association of a recombinant Cotton leaf curl Bangalore virus with yellow vein and leaf curl disease of okra in India. Indian J Virol 24:188–198

Abelmoschus manihot (Aibika, Bele)

13

Venkataravanappa V, Lakshminarayana RC, Jalali S, Krishna RM (2014) Association of Tomato leaf curl New Delhi virus DNA-B with bhendi yellow vein mosaic virus in okra showing yellow vein mosaic disease symptoms. Acta Virol 59:125–139 Venkataravanappa V, Lakshminarayana Reddy CN, Jalali S, Briddon RW, Krishna Reddy M (2015a) Molecular identification and biological characterisation of a Begomovirus associated with okra enation leaf curl disease in India. Eur J Plant Pathol 141:217–235 Venkataravanappa V, Lakshminarayana Reddy CN, Jalali S, Krishna Reddy M (2015b) Association of Tomato leaf curl New Delhi virus DNA-B with bhendi yellow vein mosaic virus in okra showing yellow vein mosaic disease symptoms. Acta Virol 59(2):125–139 Xie K, Cai JH, Hu DM, Wei X, Jia Q, Qin BX, Chen BS, Meng JR, Liu YL (2010) First report of Okra leaf curl disease in China. J Plant Pathol 92:S4.109 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zhou X (2013) Advances in understanding Begomovirus satellites. Annu Rev Phytopathol 51:357–381 Zhou X, Liu Y, Robinson DJ, Harrison BD (1998) Four DNA-A variants among Pakistani isolates of Cotton leaf curl virus and their affinities to DNA-A of geminivirus isolates from okra. J Gen Virol 79:915–923 Zia-Ur-Rehman M, Hameed U, Ali CA, Haider MS, Brown JK (2017) First report of Chickpea chlorotic dwarf virus infecting okra in Pakistan. Plant Dis 101:1336

Abelmoschus manihot (Aibika, Bele) Synonyms Hibiscus manihot Family: Malvaceae

Ornamental

Emilia yellow vein virus Taxonomic position Genus: Begomovirus

(EmYVV)

Family: Geminiviridae

EmYVV infection in plants of Abelmoschus manihot was reported from China (Tang and Qin 2011). The virus-infected aibika plants exhibit leaf curl and yellow vein symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of EmYVV, refer to Emilia sonchifolia.

Hibiscus chlorotic ringspot virus Taxonomic position Genus: Betacarmovirus

(HCRSV)

Family: Tombusviridae

HCRSV infection in plants of Abelmoschus manihot was reported from Fiji, Papua New Guinea, South Pacific Islands, Solomon Islands, Vanuatu, and the USA (Jones et al. 1998; Brunt and Spence 2000). The virus-infected aibika plants exhibit leaf chlorosis symptoms. The virus is mechanically saptransmissible. For more details of HCRSV, refer to Hibiscus rosa-sinensis.

A

14

Abrus precatorius (Jequirity)

References Brunt AA, Spence NJ (2000) The natural occurrence of Hibiscus chlorotic ringspot virus (Carmovirus; Tombusviridae) in aibika or bele (Abelmoschus manihot) in some South Pacific Island countries. Plant Pathol 49(6):798 Jones P, Devonshire J, Dabek A, Howells C (1998) First report of Hibiscus chlorotic ringspot carmovirus in Tuvalu. Plant Dis 82:591 Tang MQ, Qin LY (2011) Molecular identification of geminivirus inducing vein yellowing in Abelmoschus manihot. Zhong Yao Cai 34:168–171

Abrus precatorius (Jequirity) Family: Fabaceae

Medicinal

Clitoria yellow vein virus Taxonomic position Genus: Tymovirus

(CYVV)

Family: Tymoviridae

CYVV infection in plants of Abrus precatorius was reported from Kenya (Bock et al. 1977). The virusinfected jequirity plants exhibit yellowing of secondary and smaller leaf veins. The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of CYVV, refer to Clitoria ternatea.

References Bock KR, Guthrie EJ, Meredith G (1977) Clitoria yellow vein virus, a tymovirus from Kenya. Ann Appl Biol 85:97–103

Abutilon striatum (Redvein flowering maple) Synonyms Abutilon pictum Family: Malvaceae

Abutilon mosaic virus

Ornamental

(AbMV)

Synonyms Abutilon infectious variegation virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Abutilon striatum (Redvein flowering maple)

15

Geographical distribution AbMV infection in plants of Abutilon striatum was reported from South and Central American region, Mexico, Germany, Brazil, Argentina, Hawaii, India, and New Zealand (de Souza and Kim 1990; Wege et al. 2000; Lyttle and Guy 2004; De La Torre-Almaraz et al. 2006; Nelson 2008; Paprotka et al. 2010; Jyothsna et al. 2013). Symptoms and host(s) The virus-infected redvein flowering maple plants exhibit symptoms of heavily bright whitish- to yellow-mottled leaves, a yellow mosaic resembling variegation. The yellow patches are sharply delimited by leaf veins, giving them an angular appearance. Symptoms may vary seasonally depending on light intensity (Nelson 2008). AbMV has a very wide host range, and the hosts and reactions are as follows: Abutilon spp., Sida spp., Hibiscus spp., and Gossypium hirsutum (yellow mosaic); Malva spp. and Phaseolus vulgaris (mosaic); and Abutilon pictum (foliar crinkling, mottling, and mild mosaic). Transmission Brazilian isolates of AbMV have been shown to be transmitted by the whitefly Bemisia tabaci (Flores and Silberschmidt 1967). A single whitefly was shown to be sufficient for transmission, females being more effective than males. The minimum total acquisition and inoculation access periods were 15–20 min. The virus is transmissible by grafting but not transmissible by contact between plants. No seed and pollen transmission is observed (Costa and Carvalho 1960). Significantly, AbMV isolates that have been maintained in Abutilon by vegetative propagation for a long time are not insect (B. tabaci) transmissible (de Souza and Kim 1990; Wu et al. 1996; Hofer et al. 1997). Virion properties and genome AbMV virions were shown to be geminate (twinned quasi-icosahedra), non-enveloped, 20  29–33 nm (Abouzid and Jeske 1986). AbMV is a typical bipartite Begomovirus native to the New World. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A contains 2632 nt (X15983 = NC_001928) and DNA-B of 2585 nt (X15984 = NC_001929) (Frischmuth et al. 1990). Typical of the majority of bipartite begomoviruses native to the New World, the DNA-A component of AbMV encodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The transcripts of AbMV have been mapped and shown to be polyadenylated (Frischmuth et al. 1991). Replacement of the coat protein gene of the whitefly non-transmissible clone of AbMV with the coat protein gene of a transmissible isolate of Sida golden mosaic virus was shown to restore insect transmissibility (Hofer et al. 1997). The DNA replication of AbMV, and likely all geminiviruses, in the nuclei of host cells was shown to occur by both rolling circle replication and recombination-dependent replication (Abouzid and Jeske 1986; Groning et al. 1987; Frischmuth et al. 1990; Wu et al. 1996; Briddon 2001; Jeske et al. 2001; Paprotka et al. 2010; Jyothsna et al. 2013; Brown et al. 2015; Zerbini et al. 2017).

Abutilon yellow mosaic virus

(AbYMV)

Taxonomic position AbYMV is a tentative member of the genus Tobamovirus and family Virgaviridae.

A

16

Abutilon striatum (Redvein flowering maple)

Geographical distribution AbYMV infection in plants of Abutilon x hybrida was reported from Minnesota (Almeyda-Becerra and Lockhart 2008). Symptoms and host(s) The virus-infected redvein flowering maple plants exhibit symptoms of intense yellow leaf mottle and mosaic and reduced height, as well as loss of the characteristic red color of the flowers, which were instead pale pink. Transmission The virus has no known natural vector. The virus is mechanically sap-transmissible. Virion properties and genome The virions are rigid rods. The genome consists of a positive-sense single-stranded RNA. A partial sequence of 5093 nt is available (EU559678) (King et al. 2012; Adams et al. 2017).

References Abouzid A, Jeske H (1986) The purification and characterization of gemini particles from Abutilon mosaic virus infected Malvaceae. J Phytopathol 115:344–353 Adams MJ, Adkins S, Bragard C, Gilmer D, Li D, MacFarlane SA, Wong S-M, Melcher U, Ratti C, Ryu KH, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Virgaviridae. J Gen Virol 98:1999–2000 Almeyda-Becerra CV, Lockhart BE (2008) Characterization of Abutilon yellow mosaic virus, a tobamovirus occurring in flowering maple (Abutilon  hybrida). Phytopathology 98:S12 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Costa AS, Carvalho AMB (1960) Mechanical transmission and properties of the Abutilon mosaic virus. Phytopathol Z 37:259–272 De la Torre-Almaraz R, Romero-Rodriguez A, Monsalvo-Reyes AC, Medina-Ramos G, Torres-Pacheco I, Ruiz-Medrano R (2006) Biological characterization of a geminivirus associated with Abutilon (Abutilon x hybridum Hort. Ex. Voss. Malvacea) yellow mosaic virus in Mexico. Agrociencia 40:335–347 de VBV S, Kim KS (1990) Cytopathology and transmission characteristics of a virus isolate from Abutilon striatum. Phytopathology 80:548–552 Flores E, Silberschmidt K (1967) Contribution to the problem of insect and mechanical transmission of infectious chlorosis of Malvaceae and the disease displayed by Abutilon thompsonii. Phytopath Z 60:181–195 Frischmuth T, Zimmat G, Jeske H (1990) The nucleotide sequence of Abutilon mosaic virus reveals prokaryotic as well as eukaryotic features. Virology 178(2):461–468 Frischmuth S, Frischmuth T, Jeske H (1991) Transcript mapping of abutilon mosaic virus, a geminivirus. Virology 11815:596–604 Groning BR, Abouzid A, Jeske H (1987) Single-stranded DNA from Abutilon mosaic virus is present in the plastids of infected Abutilon sellovianum. Proc Natl Acad Sci U S A 84:8996–9000 Hofer P, Bedford ID, Markham PG, Jeske H, Frischmuth T (1997) Coat protein replacement results in whitefly transmission of an insect non transmissible geminivirus isolate. Virology 236:288–295 Jeske H, Lutgemeier M, Preiß W (2001) DNA forms indicate rolling circle and recombination-dependent replication of Abutilon mosaic virus. EMBO J 20:6158–6167 Jyothsna P, Haq QMI, Jayaprakash P, Malathi VG (2013) Molecular evidence for the occurrence of Abutilon mosaic virus, a new world Begomovirus in India. Indian J Virol 24:284–288 King AMQ, Adams MJ, Carstens EB, Lefkowitz EJ (2012) Virus taxonomy: classification and nomenclature of viruses, ninth report of the international committee on taxonomy of viruses. Elsevier Academic Press, San Diego Lyttle DJ, Guy PL (2004) First record of Geminiviruses in New Zealand: Abutilon mosaic virus and Honeysuckle yellow vein virus. Aust Plant Pathol 33:321–322

Abutilon theophrasti (Velvetleaf)

17

Nelson SC (2008) Abutilon mosaic. University of Hawaii at Manoa, College of Tropical Agriculture and Human Resources, Cooperative Extension Service, PD-39 Paprotka T, Metzler V, Jeske H (2010) The complete nucleotide sequence of a new bipartite Begomovirus from Brazil infecting Abutilon. Arch Virol 155:813–816 Wege C, Gotthardt R-D, Frischmuth T, Jeske H (2000) Fulfilling Koch’s postulates for Abutilon mosaic virus. Arch Virol 145:2217–2225 Wu ZC, Hu JS, Polston JE, Ullman DE, Hiebert E (1996) Complete nucleotide sequence of a nonvector-transmissible strain of Abutilon mosaic geminivirus in Hawaii. Phytopathology 86:608–613 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Abutilon theophrasti (Velvetleaf) Family: Malvaceae

Weed host

Abutilon golden mosaic virus Taxonomic position Genus: Begomovirus

(AbGMV)

Family: Geminiviridae

Geographical distribution AbGMV infection in plants of Abutilon theophrasti was reported from Yucatan, Mexico (TorresHerrera et al. unpublished - KC430935). Symptoms and host(s) The virus-infected velvetleaf plants exhibit bright yellow mosaic symptoms. Transmission The transmission of AbGMV has not been investigated. It is likely that, in common with other begomoviruses, the virus is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of AbGMV has not been investigated. In common with all geminiviruses, the virions of AbGMV are likely geminate (twinned icosahedra). It is unclear at this time whether AbGMV is a bipartite Begomovirus, as is typical of the majority of begomoviruses native to the New World, or monopartite since only the sequence of a single genome/ DNA-A genomic component has been submitted to the nucleotide sequence databases (KC430935). Only one monopartite Begomovirus has been confirmed to occur in the New World, tomato leaf deformation virus. The available sequence of AbGMV is a circular molecule of 2629 nt (KC430935). Typical of the DNA-A component of bipartite begomoviruses and the genomes of monopartite begomoviruses native to the New World, the sequence is predicted to encode five genes, one in the virion-sense and four in the complementary-sense. The expression and function of these genes have not been investigated for AbGMV (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

A

18

Abutilon theophrasti (Velvetleaf)

Abutilon mosaic Bolivia virus Taxonomic position Genus: Begomovirus

(AbMBoV)

Family: Geminiviridae

Geographical distribution AbMBoV infection in plants of Abutilon spp. was reported from Bolivia (Wyant et al. 2011). Symptoms and host(s) The virus-infected velvetleaf plants exhibit yellow mosaic symptoms. Transmission The transmission of AbMBoV has not been investigated. It is likely that, in common with other begomoviruses, the virus is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of AbMBoV has not been investigated. In common with all geminiviruses, the virions of AbMBoV are likely geminate (twinned icosahedra). The virus is a typical bipartite Begomovirus native to the New World. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A contains 2717 nt (HM585445 = NC_015045) and DNA-B of 2682 nt (HM585446 = NC_015048) (Briddon 2001; Wyant et al. 2011; Brown et al. 2015). Typical of the majority of bipartite begomoviruses native to the New World, the DNA-A component of AbMBoVencodes six genes, two in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for AbMBoV (Zerbini et al. 2017).

Abutilon mosaic Brazil virus Taxonomic position Genus: Begomovirus

(AbMBV)

Family: Geminiviridae

Geographical distribution AbMBV infection in plants of Abutilon theophrasti was reported from Brazil (Paprotka et al. 2010; Wyant et al. 2012). Symptoms and host(s) The virus-infected velvetleaf plants exhibit mosaic and vein-clearing symptoms. Transmission The transmission of AbMBV has not been investigated. It is likely that, in common with other begomoviruses, the virus is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of AbMBV has not been investigated. In common with all geminiviruses, the virions of AbMBV are likely geminate (twinned icosahedra).

Abutilon theophrasti (Velvetleaf)

19

AbMBV is a typical bipartite Begomovirus native to the New World. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A contains 2649 nt (JF694480 = NC_016574) and DNA-B of 2633 nt (JF694479 = NC_016577; JF694483). Typical of the majority of bipartite begomoviruses native to the New World, the DNA-A component of AbMBV encodes six genes, two in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for AbMBV (Briddon 2001; Wyant et al. 2012; Brown et al. 2015; Zerbini et al. 2017).

Abutilon yellows virus Taxonomic position Genus: Crinivirus

(AbYV)

Family: Closteroviridae

Geographical distribution AbYV infection was reported on the common weed velvetleaf (Abutilon theophrasti Medic.) was reported from Illinois, USA in 1977 (Liu et al. 1997). Symptoms and host(s) The virus-infected velvetleaf plants exhibit symptoms of extensive chlorosis to yellow discoloration of the leaves, often accompanied by stunting (Wisler and Duffus 2001). Transmission The virus is transmitted by the whitefly vector, Trialeurodes abutilonea (banded-wing whitefly), in a semi-persistent manner, and the virus is retained by the whitefly for up to 3 days (Liu et al. 1998; Jones 2003). Transmission efficiency varied from 4% for individual whiteflies to 81% for 50 whiteflies with acquisition access periods (AAP) of 24 h and inoculation access periods (IAP) of 48 h, whereas the efficiency of virus acquisition varied from 19% for single whiteflies to 77% when 50 of the insects were used (Wisler and Duffus 2001). No crop plants have been identified as hosts; however, the virus can infect members of the Malvaceae and Nicotiana clevelandii (Liu et al. 1997). The virus is not mechanically transmissible. Virion properties and genome The virions are non-enveloped, bipartite filamentous particles about 650–850 nm and 700–900 nm in length, and 10–13 nm in diameter (Liu et al. 1997, 2000), but the genome remains uncharacterized, with the exception of the coat protein and a region of the replication-associated polyprotein genes (AY422609, AY422070) (Kreuze 2011; Tzanetakis et al. 2013).

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV-infected Abutilon theophrasti plants were reported from China (Niu et al. 2009). The virusinfected velvetleaf plants exhibit symptoms of diffuse chlorotic and necrotic spots on the oldest leaves. No definite insect is identified as vector. The virus is mechanically sap-transmissible. The virus also spreads through mechanical transmission by field workers. For more details of TMV, refer to Nicotiana tabacum.

A

20

Turnip mosaic virus Taxonomic position Genus: Potyvirus

Acalypha australis (Asian copperleaf)

(TuMV)

Family: Potyviridae

TuMV infection in plants of Abutilon theophrasti was reported from Italy (Guglielmone et al. 2000). The virus-infected velvetleaf plants exhibit severe mosaic symptoms and a reduction in growth but without symptoms on flowers. The virus is easily transmitted by aphid vectors, Aphis gossypii and Myzus persicae, in a non-persistent manner, and also by mechanical sap-inoculation to a large number of host plants. The virus is not seed-transmitted in A. theophrasti. For more details of TuMV, refer to Brassica rapa.

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Guglielmone L, Jenner CE, Walsh JA, Ramasso E, Marian D, Roggero P (2000) An unusual isolate of Turnip mosaic potyvirus from Abutilon theophrasti in Piedmont, Italy. Phytoparasitica 28(2):149–152 Jones DR (2003) Plant viruses transmitted by whiteflies. Eur J Plant Pathol 109:195–219 Kreuze JF (2011) Crinivirus. Closteroviridae. In: The Springer index of viruses. Springer, New York, pp 335–342. https:// doi.org/10.1007/978-0-387-95919-1_51 Liu H-Y, Li RH, Wisler GC, Duffus JE (1997) Characterization of Abutilon yellows virus – a new clostero-like virus transmitted by banded-wing whitefly (Trialeurodes abutilonea). Phytopathology 87:S58–S59 Liu HY, Wisler GC, Duffus JE (1998) Abutilon yellow virus – a new Closterovirus transmitted by banded-wing whitefly (Trialeurodes abutilonea). Abstracts of papers presented at the 7th international congress of plant pathology, Edinburgh, GB, 1998-08-09/16 (Abst. 1.11.8) Liu H-Y, Wisler GC, Duffus JE (2000) Particle length of whitefly-transmitted criniviruses. Plant Dis 84:803–805 Niu YB, Qing L, Yao M, Wang DF, Liu JD, Wang JS (2009) First report of Tobacco mosaic virus in Abutilon theophrasti in China. Plant Dis 93:1221 Paprotka T, Metzler V, Jeske H (2010) The complete nucleotide sequence of a new bipartite begomovirus from Brazil infecting Abutilon. Arch Virol 155(5):813–816 Wisler GC, Duffus JE (2001) Transmission properties of whitefly-borne criniviruses and their impact on virus epidemiology. In: Harris KF, Smith OP, Duffus JE (eds) Virus-insect-plant interactions. Academic, New York, pp 293–306 Wyant PS, Gotthardt D, Schafer B, Krenz B, Jeske H (2011) The genomes of four novel begomoviruses and a new Sida micrantha mosaic virus strain from Bolivian weeds. Arch Virol 156(2):347–352 Wyant PS, Strohmeier S, Schafer B, Krenz B, Assuncao IP, Lima GS, Jeske H (2012) Circular DNA genomics (circomics) exemplified for geminiviruses in bean crops and weeds of northeastern Brazil. Virology 427(2):151–157 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Acalypha australis (Asian copperleaf) Family: Euphorbiaceae

Medicinal plant

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

Achimenes spp.

21

TYLCV infection in plants of Acalypha australis was reported from China and South Florida (Ying and Davis 2000; Ji et al. 2013). The virus-infected Asian copperleaf plants do not show any symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

References Ji YH, Zhang H, Zhang K, Li G, Lian S, Cheng ZB, Zhou YJ (2013) First report of Tomato yellow leaf curl virus in Acalypha australis in China. Plant Dis 97:430–431 Ying ZT, Davis MJ (2000) Partial characterization and host range of Tomato yellow leaf curl virus in South Florida. Proc Florida State Hortic Soc 113:185–190

Acanthospermum hispidum (Bristly starbur) Family: Asteraceae

Weed host

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Acanthospermum hispidum was reported from Georgia (USA) (Mullis et al. 2009). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting, but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Mullis S, Bertrand PF, Brown SL, Csinos A, Diaz-Perez JC, Gitaitis R, Hickman LL, Johnson A, LaHue SS, Martinez N, McPherson RM, Nischwitz C, Reay-Jones F, Riley D, Sherwood J, Stevenson K, Wells L (2009) Tospoviruses in Solanaceae and other crops in the coastal plain of Georgia. University of Georgia, College of Agricultural and Environmental Sciences, Bulletin 1354, pp 51

Achimenes spp. Family: Gesneriaceae

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

Ornamental

(TMV)

Family: Virgaviridae

A

22

Achyranthes bidentata (Ox knee)

TMV infection in plants of Achimenes spp. was reported from Germany and the USA (Koenig and Lesemann 1973; Zettler and Nagel 1983). The virus-infected achimenes plants exhibit symptoms of leaf mottling, narrow malformed leaves, and stunting of plants. There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

References Koenig R, Lesemann DE (1973) Tabakmosaikvirus in Achimenes. Phytopath Z 76:87–89 Zettler FW, Nagel J (1983) Infection of cultivated gesneriads by two strains of Tobacco mosaic virus. Plant Dis 67:1123–1125

Achyranthes bidentata (Ox knee) Family: Amaranthaceae

Medicinal plant

Alternanthera mosaic virus Taxonomic position Genus: Potexvirus

(AltMV)

Family: Alphaflexiviridae

AltMV infection in plants of Achyranthes bidentata was reported from the Asia (Iwabuchi et al. 2016). The virus-infected Ox knee plants exhibit mosaic symptoms on the leaves. No vector is reported for this virus. The virus is readily sap-transmissible and easily spread by plant contact, vegetative propagation, and contaminated tools and hands. For more details of AltMV, refer to Alternanthera spp.

References Iwabuchi N, Yoshida T, Yusa A, Nishida S, Tanno K, Keima T, Nijo T, Yamaji Y, Namba S (2016) Complete genome sequence of Alternanthera mosaic virus, isolated from Achyranthes bidentata in Asia. Genome Announc 4: e00020–e00016

Aconitum spp. Family: Ranunculaceae

Aconitum latent virus Taxonomic position Genus: Carlavirus

Ornamental

(AcLV)

Family: Betaflexiviridae

Aconitum spp.

23

Geographical distribution AcLV infection in plants of Aconitum napellus was first reported from Israel (Cohen et al. 2000). The virus spreads in Israel and Japan. Symptoms and host(s) The virus-infected aconitum plants do not exhibit any symptoms. Transmission The virus is transmitted by an aphid vector Myzus persicae in a non-persistent manner, and also through vegetative propagation. Virion properties and genome The virions are flexuous filaments about 640 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8657 nt (AB051848 = NC_002795) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Adams et al. 2004).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Aconitum spp. was reported from Israel and Japan (Cohen et al. 1998; Fukumoto et al. 2008). The virus-infected aconitum plants display mild mosaic symptoms. The virus is transmitted by aphid vectors Myzus persicae, Aphis gossypii, Acrythosiphon pisum, and Aphis craccivora in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Helleborus mosaic virus Taxonomic position Genus: Carlavirus

(HeMV)

Family: Betaflexiviridae

HeMV infection in plants of Aconitum spp. was reported from Germany (Richert-Poggeler et al. 2015). The virus-infected aconitum plants exhibit symptoms of chlorotic and/or necrotic lesions along the leaf veins. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of HeMV, refer to Helleborus spp.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Cohen J, Zeidan M, Franck A, Bekelman E, Gotman S, Gera A (1998) Viruses in Aconitum. Phytoparasitica 26:166–167

A

24

Actinidia chinensis (Kiwifruit)

Cohen J, Zeidan M, Rosner A, Gera A (2000) Biological and molecular characterization of a new carlavirus isolated from an Aconitum sp. Phytopathology 90:340–344 Fukumoto F, Fuji SI, Shinoda K, Iizuka M (2008) Cucumber mosaic virus isolated from Aconitum spp. in Japan. J Gen Plant Pathol 74:88–90 Richert-Poggeler KR, Turhal A-K, Schuhmann S, Maa C, Blockus S, Zimmermann E, Eastwell KC, Martin RR, Lockhart B (2015) Carlavirus biodiversity in horticultural host plants: efficient virus detection and identification combining electron microscopy and molecular biology tools. Acta Hortic 1072:37–45

Actinidia chinensis (Kiwifruit) Family: Actinidiaceae

Edible fruit

Actinidia chlorotic ringspot-associated emaravirus Taxonomic position Genus: Emaravirus

(AcCRaV)

Family: Fimoviridae

Geographical distribution AcCRaV infection in plants of Actinidia chinensis was reported from China (Zheng et al. 2017). Symptoms and host(s) The virus-infected kiwifruit plants exhibit chlorotic spots, chlorotic ring-spots, and vein-yellowing symptoms on leaves. Transmission The virus is transmitted by an eriophyid mite vector. The virus is also mechanically sap-transmissible to Nicotiana benthamiana on which chlorotic and ring-spots develop. Virion properties and genome The virions are spherical, enveloped by a double membrane, and measure from 80 to 100 nm in diameter. The genome consists of a multipartite negative-sense, single-stranded RNA genome with five genomic components: RNA1 7061 nt (KT861481), RNA2 2267 nt (KT861482), RNA3 6078 nt (KT861483), RNA4 1664 nt (KT861484), and RNA5 1474 nt (KT861485) (Zheng et al. 2017; Elbeaino et al. 2018).

Actinidia seed-borne latent virus Taxonomic position Genus: Prunevirus

(ASbLV)

Family: Betaflexiviridae

Geographical distribution ASbLV infection in plants of Actinidia chinensis was reported from Auckland (New Zealand) and South Korea (Veerakone et al. 2018; Park et al. unpublished – LC438404)

Actinidia chinensis (Kiwifruit)

25

Symptoms and host(s) Initially reported plants of A. chinensis raised from seed showed symptoms of veinal chlorosis and mottling, but were co-infected with Actinidia virus A; additional plants with a single infection of ASbLV were symptomless. No other hosts are reported. Transmission The virus is naturally transmitted through seed, and is also mechanically sap-transmissible. The use of infected bud wood may be the primary source of virus spread. Virion properties and genome The virions are flexuous filaments, usually 12–13 nm in diameter (range 10–15 nm) and from 600 to over 1000 nm in length. The genome consists of a single molecule of linear single-stranded RNA of 8192 nt (MF440375 = NC_040800; LC438404) excluding the 30 poly(A) tail, and contains four open reading frames (Veerakone et al. 2018).

Actinidia virus 1 Taxonomic position Genus: Unassigned

(AcV-1)

Family: Closteroviridae

Geographical distribution AcV-1 infection in plants of Actinidia chinensis was reported from Italy and China (Blouin et al. 2018; Peng et al. 2019). Symptoms and host range The virus-infected kiwifruit plants exhibit leaf mottle, chlorosis, and malformation symptoms. Transmission The virus is transmitted by mechanical sap inoculation. Virion properties and genome The virions are filaments usually flexuous with a clear modal length of 1717 nm. The genome is a monopartite, linear, positive-sense, single-stranded RNA of 18848 nt (KX857665 = NC_035453) and contains 12 open reading frames (ORFs) greater than 6 KDa, one carrying two papain-like leader proteases, a methyltransferase, a helicase, and an RNA-dependent RNA polymerase domain. Additional ORFs code for homologs of heat shock protein 70, heat shock protein 90, and a coat protein.

Actinidia virus A Taxonomic position Genus: Vitivirus

(AcVA)

Family: Betaflexiviridae

Geographical distribution AcVA infection in plants of Actinidia chinensis was reported from New Zealand, Italy, South Korea, and China (Blouin et al. 2012, 2013; Zheng et al. 2014; Cho et al. 2017).

A

26

Actinidia chinensis (Kiwifruit)

Symptoms and host(s) Symptoms observed on leaves of A. chinensis included leaf vein chlorosis, flecking, and ring-spots. No symptoms were detected in A. deliciosa. Symptoms were seen most commonly on basal leaves of new shoots around flowering time. Symptoms on some veins faded and were difficult to observe in summer, while symptoms on other veins did not appear until early summer. Transmission The virus is mechanically sap-transmissible to Nicotiana occidentalis Virion properties and genome The virions are helically constructed flexuous filaments 750–800  12 nm, showing distinct crossbanding. The genome consists of a single molecule of positive-sense ssRNA of 7566 nt (JN427014) and contains five slightly overlapping ORFs which encode, in order, the replication-related proteins, a 19–20 K protein with unknown functions, a MP of the “30 K” superfamily type, the CP, and a small protein (10–14 K) with nucleotide-binding properties. Coat protein subunits are of one type and 18–21.5 kDa in size (Adams et al. 2004; Blouin et al. 2012).

Actinidia virus B Taxonomic position Genus: Vitivirus

(AcVB)

Family: Betaflexiviridae

Geographical distribution AcVB infection in plants of Actinidia chinensis was reported from New Zealand, Italy, South Korea, and China (Blouin et al. 2012, 2013; Zheng et al. 2014; Cho et al. 2017). Symptoms and host(s) The virus-infected A. chinensis plants exhibit symptoms of vein chlorosis, flecking, and ring-spots on the leaves. No symptoms were detected in A. deliciosa. Symptoms were seen most commonly on basal leaves of new shoots around flowering time. Symptoms on some veins faded and were difficult to observe in summer, while symptoms on other veins did not appear until early summer. Transmission The virus is mechanically sap-transmissible to Nicotiana occidentalis Virion properties and genome The virions are helically constructed flexuous filaments 750–800  12 nm, showing distinct crossbanding. The genome consists of a single molecule of positive-sense ssRNA of 7488 nt (JN427015 = NC_016404) and contains five slightly overlapping ORFs which encode, in order, the replicationrelated proteins, a 19–20 K protein with unknown functions, a MP of the “30 K” superfamily type, the CP, and a small protein (10–14 K) with nucleotide-binding properties. Coat protein subunits are of one type and 18–21.5 kDa in size (Adams et al. 2004; Blouin et al. 2012).

Actinidia chinensis (Kiwifruit)

Actinidia virus X Taxonomic position Genus: Potexvirus

27

(AVX)

Family: Alphaflexiviridae

Geographical distribution AVX infection in plants of Actinidia chinensis was reported from New Zealand (Pearson et al. 2011). Symptoms and host(s) The virus-infected kiwifruit plants do not exhibit any symptoms. Transmission The virus is transmitted by means not involving a vector. The virus transmitted by mechanical sapinoculation. Use of infected planting material is the primary source of virus spread. Virion properties and genome The virions are flexuous filaments about 485 nm long and 12–13 nm wide. The genome is monopartite positive-sense single-stranded RNA of 6888 nt (KR872420 = NC_028649) and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type, 18–27 kDa in size (James and Phelan 2016).

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

(AMV)

Family: Bromoviridae

AMV infection in plants of Actinidia chinensis was reported from New Zealand (Pearson et al. 2011; Blouin et al. 2013). The virus-infected kiwifruit plants exhibit yellow mosaic symptoms on the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of AMV, refer to Medicago sativa.

Apple stem grooving virus Taxonomic position Genus: Capillovirus

(ASGV)

Family: Betaflexiviridae

ASGV has been identified in plants of Actinidia chinensis was reported from New Zealand, China, and India (Clover et al. 2003; Pearson et al. 2010; Blouin et al. 2013). The leaves of virus-infected kiwifruit plants exhibit a variety of symptoms, including interveinal mottling, chlorotic mosaics, and ring-spots. No vector has been identified for this virus. The virus is mechanically sap-transmissible. The virus is transmitted by grafting and disseminated by infected propagative material. For more details of ASGV, refer to Malus domestica.

A

28

Actinidia chinensis (Kiwifruit)

Cherry leaf roll virus

(CLRV)

Taxonomic position Genus: Nepovirus

Family: Secoviridae

CLRV infection in plants of Actinidia chinensis was reported from New Zealand (Pearson et al. 2010; Woo et al. 2012; Blouin et al. 2013). The virus-infected kiwifruit plants exhibit symptoms of leaf spots, bark cracking, cane wilting, fruit malformation, and reduced yields. The virus is transmitted by nematode vectors, Xiphinema coxi, X. diversicaudatum, and X. vuittenezi, in a non-persistent manner, and also by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants. For more details of CLRV, refer to Prunus avium.

Citrus leaf blotch virus Taxonomic position Genus: Citrivirus

(CLBV)

Family: Betaflexiviridae

CLBV infection in plants of Actinidia chinensis was reported from New Zealand and China (Pearson et al. 2010; Blouin et al. 2013; Chavan et al. 2013). Symptoms of virus-infected Actinidia, vein-clearing and mild mottling, were observed on the leaves of the terminal and middle regions of shoots of the infected A. chinensis male plant (M3-A) during spring and interveinal chlorosis during summer. Similar symptoms were observed in female A. chinensis accessions (F1-A and F3-N8), while five other accessions (F1-N, F3-E4, F3-P, F3-QII, and F4-J) remained symptomless for 4 years. No vector is reported for this virus. The virus is mechanically sap-transmissible to N. benthamiana, N. clevelandii, N. occidentalis 37B, and N. glutinosa. The primary spread of the virus takes place by the use of infected bud sticks. For more details of CLBV, refer to Citrus spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Actinidia chinensis was reported from Italy and New Zealand (Pearson et al. 2010, 2011; Blouin et al. 2013). The virus-infected kiwifruit plants exhibit chlorosis symptoms on the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Cucumber necrosis virus Taxonomic position Genus: Tombusvirus

(CNV)

Family: Tombusviridae

CNV infection in plants of Actinidia chinensis was reported from China, Italy, and New Zealand (Pearson et al. 2011; Blouin et al. 2013). The virus-infected kiwifruit plants do not exhibit any symptoms. The virus is transmitted by a fungal vector, Olpidium radicale, and also by mechanical sap-inoculation. The virus is not transmitted by seed or pollen. For more details of CNV, refer to Cucumis sativus.

Actinidia chinensis (Kiwifruit)

29

Pelargonium zonate spot virus Taxonomic position Genus: Anulavirus

(PZSV)

Family: Bromoviridae

PZSV infection in plants of Actinidia chinensis was reported from Italy, Spain, France, Israel, the USA, and New Zealand (Biccheri et al. 2012; Blouin et al. 2013). The virus-infected kiwifruit plants exhibit symptoms including chlorotic and necrotic rings on the leaves and depressed areas on the fruits with consequent deformation of the berries. Thrips are involved in transmitting this virus by carrying the virus-infected pollen grains on their bodies and introduce the virus into the susceptible hosts while feeding. The virus is mechanically sap-transmissible to Chenopodium quinoa, Nicotiana benthamiana, N. glutinosa, and N. tabacum. For more details of PZSV, refer to Pelargonium spp.

Ribgrass mosaic virus Taxonomic position Genus: Tobamovirus

(RMV)

Family: Virgaviridae

RMV infection in plants of Actinidia chinensis was reported from China and New Zealand (Pearson et al. 2007, 2010; Chavan et al. 2012; Blouin et al. 2013). The virus-infected A. chinensis accessions exhibit chlorosis of the leaf veins and adjacent tissue at the tip and middle regions of shoots during spring, followed by chlorotic mottles, mosaics, and ring-spots during summer. Some leaves with vein chlorosis also displayed mild hypertrophy of the mesophyll tissue. Symptoms on A. deliciosa plants included chlorotic mottling or mosaic during spring and ring-spots during the summer months. No vector is reported for this virus. The virus is mechanically sap-transmissible. The virus is transmissible by grafting and also by contact between plants. For more details of RMV, refer to Plantago lanceolata.

Turnip vein-clearing virus Taxonomic position Genus: Tobamovirus

(TVCV)

Family: Virgaviridae

TVCV infection in plants of Actidinia chinensis was reported from China and New Zealand (Chavan et al. 2009; Blouin et al. 2013). The virus-infected kiwifruit plants do not exhibit any symptoms. The virus is mechanically sap-transmissible and also by contact between plants. No vector is known for this virus. For more details of TVCV, refer to Brassica rapa.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Biccheri R, Babini AR, Blouin A, Lanzoni C, Pisi A, Poggi Pollini C, Credi R, Laghi L, Rocculi P, Rubies Autonell C, Pearson MN, Ratti C (2012) Pelargonium zonate spot virus infecting kiwifruit plants in Italy. Abstract of a paper presented at the 22nd international conference on virus and other graft transmissible diseases of fruit crops (Rome, 2012-06-03/08), p 27

A

30

Actinidia deliciosa (Fuzzy kiwifruit)

Blouin AG, Chavan RR, Pearson MN, MacDiarmid RM, Cohen D (2012) Detection and characterisation of two novel vitiviruses infecting Actinidia. Arch Virol 157:713–722 Blouin AG, Pearson MN, Chavan RR, Woo ENY, Lebas BSM, Veerakone S, Ratti C, Biccheri R, MacDiarmid RM, Cohen D (2013) Viruses of kiwifruit (Actinidia species). J Phytopathol 95:221–235 Blouin AG, Biccheri R, Khalifa ME, Pearson MN, Poggi Pollini C, Hamiaux C, Cohen D, Ratti C (2018) Characterization of Actinidia virus 1, a new member of the family Closteroviridae encoding a thaumatin-like protein. Arch Virol 163(1):229–234 Chavan RR, Pearson MN, Cohen D (2009) Partial characterization of a novel Tobamovirus infecting Actinidia chinensis and A. deliciosa (Actinidiaceae) from China. Eur J Plant Pathol 124:247–259 Chavan RR, Cohen D, Blouin AG, Pearson MN (2012) Characterization of the complete genome of Ribgrass mosaic virus isolated from Plantago major L. from New Zealand and Actinidia spp. from China. Arch Virol 157:1253–1260 Chavan RR, Blouin AG, Cohen D, Pearson MN (2013) Characterization of the complete genome of a novel Citrivirus infecting Actinidia chinensis. Arch Virol 158:1679–1686 Cho S-Y, Kim H, Yi S-I (2017) First report of Actinidia virus A and B infecting Actinidia chinensis in South Korea. Plant Dis 101:1560 Clover GRG, Pearson MN, Elliott DR, Tang Z, Smales TE, Alexander BJR (2003) Characterization of a strain of Apple stem grooving virus in Actinidia chinensis from China. Plant Pathol 52:371–378 Elbeaino T, Digiaro M, Mielke-Ehret N, Muelbach H-P, Martelli GP, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Fimoviridae. J Gen Virol 99:1478–1479 James D, Phelan J (2016) Complete genome sequence of a strain of Actinidia virus X detected in Ribes nigrum cv. Baldwin showing unusual symptoms. Arch Virol 161:507–511 Pearson MN, Chavan RR, Cohen D (2007) Viruses of Actinidia: do they pose a threat to kiwifruit production? Acta Hortic 753:639–644 Pearson MN, Cohen D, Chavan R, Blouin AG, Cowell SJ (2010) Molecular characterisation of viruses from Kiwifruit. In: 21st international conference on virus and other graft transmissible diseases of fruit crops, Neustadt. Julius-KuhnArchiv 427:87–91 Pearson MN, Cohen D, Chavan R, Blouin A (2011) Actinidia is a natural host to a wide range of plant viruses. Acta Hortic 913:467–471 Peng QD, Lv R, Ning JC, Yang T, Lin HH, Xi DH, Zhuang QG (2019) First report of Actinidia virus 1 infecting Actinidia chinensis in China. Plant Dis 103(4):782 Veerakone S, Liefting LW, Tang J, Ward LI (2018) The complete nucleotide sequence and genome organisation of a novel member of the family Betaflexiviridae from Actinidia chinensis. Arch Virol 163(5):1367–1370 Woo ENY, Lebas B, Veerakone S, Tang J, Ward L, Pearson MN (2012) Molecular detection and characterisation of Cherry leaf roll virus in New Zealand. In: 10th Australasian plant virology workshop, Hamner Springs, p 48 Zheng YZ, Wang GP, Hong N, Zhou JF, Yang ZK (2014) First report of Actinidia virus A and Actinidia virus B on Kiwifruit in China. Plant Dis 98:1590 Zheng Y, Navarro B, Wang G, Wang Y, Yang Z, Xu W, Zhu C, Wang L, Serio FD, Hong N (2017) Actinidia chlorotic ringspot-associated virus: a novel emaravirus infecting kiwifruit plants. Mol Plant Pathol 18(4):569–581

Actinidia deliciosa (Fuzzy kiwifruit) Family: Actinidiaceae

Edible fruit

Apple stem grooving virus Taxonomic position Genus: Capillovirus

(ASGV)

Family: Betaflexiviridae

ASGV infection in plants of Actinidia deliciosa was reported from India (Bhardwaj et al. 2014). The virus-infected fuzzy kiwifruit plants exhibit symptoms of severe interveinal mottling, leaf distortion, ring-spots, and chlorosis along the leaf margins. No vector has been identified for this virus. The virus is mechanically sap-transmissible. The virus is transmitted by grafting and disseminated by infected propagative material. For more details of ASGV, refer to Malus domestica.

Adenium obesum (Desert rose)

31

References Bhardwaj P, Ram R, Zaidi AA, Hallan V (2014) Characterization of Apple stem grooving virus infecting Actinidia deliciosa (Kiwi) in India. Sci Hortic 176:105–111

Adenium obesum (Desert rose) Family: Apocynaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Adenium obesum was reported from Florida, USA and Taiwan (Baker et al. 2003; Chang et al. 2012; Chen et al. 2012). The virus-infected desert rose plants exhibit symptoms of mosaic, chlorotic ring-spot, and line patterns on leaves. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Adenium obesum was reported from the Netherlands, Bulgaria, and Florida (USA) (Adkins and Baker 2005; Dikova 2011). The virus-infected desert rose plants exhibited foliar symptoms, including chlorotic ring and line pattern symptoms. The virus is transmitted by thrips vectors in a persistent, propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting, but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Adkins S, Baker CA (2005) Tomato spotted wilt virus identified in desert rose in Florida. Plant Dis 89:526 Baker CA, Achor D, Adkins S (2003) Cucumber mosaic virus diagnosed in desert rose in Florida. Plant Dis 87:1007 Chang YS, Lin YW, Chen TC, Chen YK (2012) Cucumber mosaic virus-associated ringspot mosaic disease of desert rose. Plant Pathol Bull 21:39–45 Chen YK, Chang YS, Lin YW, Wu MY (2012) First report of Cucumber mosaic virus in Desert rose in Taiwan. Plant Dis 96:593 Dikova B (2011) Tomato spotted wilt virus on some medicinal and essential oil-bearing plants in Bulgaria. Bulg J Agric Sci 17:306–313

A

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Adiantum pedatum (Fern)

Adiantum pedatum (Fern) Family: Pteridaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Adiantum pedatum was reported from the USA (Nameth and Steininger 1997). The virus-infected fern plant leaves were slightly malformed such that the pinnules were arranged irregularly at the pinna. The sequence of the pinnae on the rachis and the general appearance of the rachis were distorted. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

References Nameth ST, Steininger J (1997) Cucumber mosaic virus associated with leaf malformation and stunting in Adiantum pedatum. Plant Dis 81:1333

Aeonium spp. Family: Crassulaceae

Aeonium ringspot virus Taxonomic position Genus: Nepovirus

Ornamental

(AeRSV)

Family: Secoviridae

Geographical distribution AeRSV infection in plants of Aeonium spp. was reported from Italy (Sorrentino et al. 2013). Symptoms and host(s) The virus-infected aeonium plants exhibit chlorotic spots and rings on both leaf surfaces. Transmission The virus is mechanically sap-transmissible. Virion properties and genome The virions are isometric, non-enveloped of two types, but are similar in size, 25–30 nm in diameter, and exhibit icosahedral symmetry (T=1, pseudo T=3). The genome is a bipartite, linear positive-sense,

Aeschynanthus spp.

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single-stranded RNA. RNA1 consists of 7549 nt (JX304792) and RNA2 of 4010 nt (JQ670669). A third RNA molecule of 3472 nt entirely derived from RNA2 was also found (Sanfacon et al. 2009; Dunez and Le Gall 2011; Sorrentino et al. 2013; Sanfacon 2015; Thompson et al. 2017).

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Aeonium spp. was reported from southern Italy (Sorrentino et al. 2012). The virus-infected aeonium plants exhibit symptoms of chlorotic spots and rings on both leaf surfaces. The virus is transmitted through the nematode vector Xiphinema americanum in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

References Dunez J, Le Gall O (2011) Nepovirus. Comoviridae. In: The Springer index of viruses. Springer, New York, pp 361–369. https://doi.org/10.1007/978-0-387-95919-1_55 Sanfacon H (2015) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. In: eLS. Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000764.pub3. http://www.els.net Sanfacon H, Wellink J, Le Gall O, Karasev A, van der Vlugt R, Wetzel T (2009) Secoviridae: a proposed family of plant viruses within the order Picornavirales that combines the families Sequiviridae and Comoviridae, the unassigned genera Cheravirus and Sadwavirus, and the proposed genus Torradovirus. Arch Virol 154(5):899–907 Sorrentino R, Alioto D, Russo M, Rubino L (2012) Presence of Tobacco ringspot virus in aeonium. J Plant Pathol 94: S4.103 Sorrentino R, De Stradis A, Russo M, Alioto D, Rubino L (2013) Characterization of a putative novel nepovirus from Aeonium sp. Virus Res 177:217–221 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J Gen Virol 98:529–531

Aeschynanthus spp. Family: Gesneriaceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Aeschynanthus spp. was reported from the Czech Republic (Mertelik et al. 2002). The virus-infected aeschynanthus plants exhibit symptoms of systemic chlorotic spots and patterns. The virus is transmitted by thrips vectors in a persistent, propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

A

34

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

Aesculus hippocastanum (Horse chestnut)

(TMV)

Family: Virgaviridae

TMV infection in plants of Aeschynanthus spp. was reported from Europe and the USA (Zettler and Nagel 1983; Paludan 1985). The symptoms on the foliage of infected aeschynanthus plants were inconspicuous. There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Aeschynanthus pulcher was reported from Pennsylvania (USA) (Hausbeck et al. 1992). The virus-infected aeschynanthus plants exhibit mottle mosaic, necrotic, and chlorotic ring-spot symptoms. The virus is transmitted by thrips vectors in a persistent, propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting, but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among green house ornamentals in Pennsylvania. Plant Dis 76:795–800 Mertelik J, Mokra V, Gotzova B, Gabrielova S (2002) Occurrence and identification of Impatiens necrotic spot tospovirus in the Czech Republic. Acta Hortic 568:79–83 Paludan N (1985) Inactivation of Tobacco mosaic virus in Aeschynanthus hildebrandii by means of heat treatment chemotherapy and meristem tip culture. Tidsskrift For Planteavl 89:273–278 Zettler FW, Nagel J (1983) Infection of cultivated Gesneriads by two strains of Tobacco mosaic virus. Plant Dis 67:1123–1125

Aesculus hippocastanum (Horse chestnut) Family: Sapindaceae

Medicinal

Apple mosaic virus

(ApMV)

Taxonomic position Genus: Ilarvirus

Family: Bromoviridae

ApMV infection in plants of Aesculus hippocastanum was reported from England (Sweet and Barbara 1979). The virus-infected horse chestnut plants exhibit severe yellow mosaic symptoms. No insect vector is known for this virus. The virus is mechanically sap-transmissible and by grafting. For more details of ApMV, refer to Malus domestica.

Agapanthus spp. (African lily)

35

Prunus necrotic ringspot virus Taxonomic position Genus: Ilarvirus

(PNRSV)

A

Family: Bromoviridae

PNRSV infection in trees of Aesculus hippocastanum was reported from the UK and Europe (Sweet 1976). The virus-infected horse chestnut trees exhibit yellow mosaic symptoms. The virus is transmitted by thrips vectors, the virus present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible to a large number of hosts and by grafting. Use of planting material from infected plants is the main source of virus spread. For more details of PNRSV, refer to Prunus persica.

Strawberry latent ringspot virus

(SLRSV)

Synonyms Aesculus line pattern virus Taxonomic position Genus: Unassigned

Family: Secoviridae

SLRSV infection in trees of Aesculus hippocastanum was reported from Eastern Germany (Schmelzer 1969). The virus-infected horse chestnut trees exhibit chlorotic or necrotic ring and line pattern symptoms. The virus is transmitted by a nematode vector, Xiphinema diversicaudatum. The virus is also transmitted by mechanical sap-inoculation, and by grafting. For more details of SLRSV, refer to Fragaria spp.

References Schmelzer K (1969) Das Ulmenscheckungs-Virus. Phytopathol Z 64:39–67 Sweet JB (1976) Virus diseases of some ornamental and indigenous trees and shrubs in Britain. Acta Hortic 59:83–92 Sweet JB, Barbara DJ (1979) A yellow mosaic disease of horse chestnut (Aesculus spp.) caused by Apple mosaic virus. Ann Appl Biol 92(3):335–341

Agapanthus spp. (African lily) Family: Amaryllidaceae

Ornamental

Eggplant mottled dwarf nucleorhabdovirus Taxonomic position Genus: Nucleorhabdovirus

(EMDV)

Family: Rhabdoviridae

EMDV infection in plants of Agapanthus spp. was reported from Italy (KJ082087 = NC_025389) (Zhai et al. 2014). The virus-infected African lily plants exhibit symptoms of mosaic consisting of white

36

Agapanthus spp. (African lily)

and yellow rectangular/oval to linear patches and stripes, often confluent. The virus is transmitted by the leafhopper vectors in a persistent, propagative manner, and also by mechanical sap-inoculation. For more details of EMDV, refer to Solanum melongena.

Nerine virus X

(NVX)

Synonyms Agapanthus virus X Taxonomic position Genus: Potexvirus

Family: Alphaflexiviridae

NVX infection in plants of Agapanthus spp. was reported from the Netherlands and Taiwan (Phillips and Brunt 1980; Fuji et al. 2006; Chiang et al. 2010). The virus-infected African lily plants exhibit foliar chlorotic strip and mottle symptoms. The virus is transmitted by mechanical sap-inoculation. The virus is not transmissible by contact between plants, not transmitted by seed, and not transmitted by pollen. For more details of NVX, refer to Nerine spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Agapanthus praecox ssp. orientalis was reported from Australia (Wilson et al. 2000). The virus-infected African lily plants exhibit symptoms of concentric ring and line patterns, irregular chlorotic blotches, and streaks; the infected plants were severely stunted and did not flower. The affected leaves senesced early, and subsequently produced leaves were symptomless, perhaps suggesting only local infection. The virus is transmitted by thrips vectors in a persistentpropagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is not transmissible by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Chiang FL, Chang CA, Cheng YH, Chen CC (2010) Serological and molecular identification of Nerine virus X infecting african lily (Agapanthus africanus L.) and development of its detection techniques. J Taiwan Agric Res 60:89–100 Fuji S, Shinoda K, Furuya H, Naito H, Fukumoto F (2006) Complete nucleotide sequence of Nerine virus X (NVX-J) isolated from the African lily plant (Agapanthus campanulatus L.) in Japan. Arch Virol 151:205–208 Phillips S, Brunt AA (1980) Some hosts and properties of an isolate of Nerine virus X from Agapanthus praecox ssp. orientalis. Acta Hortic 110:65–70 Wilson CR, Wilson AJ, Pethybridge SJ (2000) First report of Tomato spotted wilt virus in common Agapanthus. Plant Dis 84:491 Zhai Y, Miglino R, Sorrentino R, Masenga V, Alioto D, Pappu HR (2014) First report of natural infection of Agapanthus sp. by Eggplant mottled dwarf virus (EMDV). New Dis Rep 29:20

Agave demeesteriana

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Agastache spp. (Korean mint) Family: Lamiaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Medicinal plant

(CMV)

Family: Bromoviridae

CMV infection in plants of Agastache spp. was reported from China and Italy (Bruni et al. 2007; Du et al. 2014). The virus-infected Korean mint plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

References Bruni R, Bianchi A, Bellardi MG (2007) Essential oil composition of Agastache anethiodora Britton (Lamiaceae) infected by Cucumber mosaic virus (CMV). Flavour Fragr J 22(1):67–70 Du J, Zhang CM, Niu YB (2014) First report of Cucumber mosaic virus in Agastache rugosa in China. Plant Dis 98:1589.3

Agave demeesteriana Family: Asparagaceae

Ornamental/succulent plant

Citrus chlorotic spot dichorhavirus (CiCSV) Taxonomic position Genus: Dichorhavirus

Family: Rhabdoviridae

CiCSV infection in plants of Agave desmettiana was reported from Brazil (Chabi-Jesus et al. 2019). The virus is transmitted by a mite vector, Brevipalpus yothersi in a persistent-propagative manner. For more details of CiCSV, refer to Citrus spp.

References Chabi-Jesus C, Ramos-González PL, Tassi AD, Barguil BM, Beserra Junior JEA, Harakava R, Kitajima EW, FreitasAstúa J (2019) First report of citrus chlorotic spot virus infecting the succulent plant Agave desmettiana. Plant Dis. https://doi.org/10.1094/PDIS-09-18-1617-PDN

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Ageratum spp. (White weed)

Ageratum spp. (White weed) Family: Asteraceae

Weed host

Ageratum enation virus Taxonomic position Genus: Begomovirus

(AEV)

Family: Geminiviridae

Geographical distribution AEV infection in plants of Ageratum conyzoides was reported from Nepal, India, and Pakistan (Kumar et al. 2011; Pandey et al. 2011; Khan et al. 2012, 2014; Srivastava et al. 2013, 2015; Tahir et al. 2015). Symptoms and host(s) The virus-infected white weed plants exhibit foliar vein yellowing which develops into a bright yellow mosaic. In some cases, infection of Ageratum is associated not with yellowing but with vein swelling and enations. However, care needs to be taken in interpreting symptoms since these are usually determined by the betasatellite with which the virus associates. The virus has been isolated from Ageratum conyzoides, Amaranthus cruentus, carrot (Daucus carota), Cleome gynandra, Crassocephalum crepidioides, and marigold (Tagetes patula). Transmission The transmission of AEV has not been investigated. It is likely that, in common with other begomoviruses, AEV will be transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Similarly, it is likely that the virus will prove not to be either mechanically or seed-transmitted. Under experimental conditions, AEV infects Nicotiana benthamiana, N. tabacum, and Solanum lycopersicum (Tahir et al. 2015). Virion properties and genome The structure of the virions of AEV has not been investigated. In common with all geminiviruses, the virions of AEV are likely geminate (twinned icosahedra). The genome of AEV consists of a single, circular single-stranded DNA of ~2746 nt (AJ437618 = NC_003434). Ageratum yellow leaf curl betasatellite (AYLCB) is required for efficient virus infection in carrot plants (Kumar et al. 2013). The genome of AEV encodes the six genes typically encoded by monopartite begomoviruses although the expression and function of these have not been investigated (Briddon 2001; Srivastava et al. 2013, 2015; Brown et al. 2015; Zerbini et al. 2017).

Ageratum latent virus Taxonomic position Genus: Ilarvirus

(AgLV)

Family: Bromoviridae

Geographical distribution AgLV infection in plants of Ageratum houstonianum was reported from Australia (Sharman and Thomas 2013).

Ageratum spp. (White weed)

39

Transmission The virus is transmitted by different thrips species. Infected pollen is blown to healthy plants where thrips feeding on the pollen mechanically transmit the virus. The virus is carried both inside the pollen and on the surface. The virus is transmitted by mechanical sap-inoculation, and by grafting. The virus is not transmitted by contact between plants. Virion properties and genome The virus has nonenveloped isometric and quasi-isometric (bacilliform) particles of 23, 25, and 27 nm. The genome is segmented, tripartite linear positive-sense ssRNA composed of RNA1 (3481 nt; JX463340 = NC_022127); RNA 2 (2872 nt; JX463341 = NC_022128); and RNA 3 (2224 nt; JX463342 = NC_022129) (Scott 2011; Sharman and Thomas 2013).

Ageratum leaf curl virus Taxonomic position Genus: Begomovirus

(ALCuV)

Family: Geminiviridae

Geographical distribution ALCuV infection in plants of Ageratum conyzoides was reported from China (Huang and Zhou 2006). Symptoms and host(s) The virus-infected white weed plants exhibit vein thickening, vein swelling, and leaf curling symptoms. The virus has been isolated from Ageratum conyzoides and Calotropis procera. Transmission The transmission of ALCuV has not been investigated. It is likely that, in common with other begomoviruses, ALCuV will be transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Similarly, it is likely that the virus will prove not to be either mechanically or seed transmitted. Virion properties and genome The structure of the virions of ALCuV has not been investigated. In common with all geminiviruses the virions of ALCuV will likely be geminate (twinned icosahedra). With only two isolates of ALCuV having been characterized, the precise nature of the genome (whether monopartite, monopartite betasatellite-associated, or bipartite) is unclear. The genome of ALCuV are each 2735 nt (AJ851005 = NC_006384; KU376491) (Briddon 2001; Huang and Zhou 2006; Brown et al. 2015; Zerbini et al. 2017). Ageratum leaf curl betasatellite DNA molecule is associated with ALCuV, and consists of 1335 nt (NC_020889) (Leke et al. 2012; Zhou 2013).

Ageratum yellow vein Hualian virus Taxonomic position Genus: Begomovirus

(AYVHuV)

Family: Geminiviridae

Geographical distribution AYVHuV infection in plants of Ageratum spp. was reported from Taiwan (Tsai et al. 2011).

A

40

Ageratum spp. (White weed)

Symptoms and host(s) The virus-infected white weed plants exhibit yellow vein mosaic symptoms. The virus was also detected in tomato. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 28  30 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2756 nt (DQ866124 = NC_010812, DQ866132) (Briddon 2001; Tsai et al. 2011; Brown et al. 2015; Zerbini et al. 2017).

Ageratum yellow vein Sri Lanka virus Taxonomic position Genus: Begomovirus

(AYVSLV)

Family: Geminiviridae

Geographical distribution AYVSLV infection in plants of Ageratum spp. was reported from Sri Lanka (Tsai et al. unpublished AF314144). Symptoms and host(s) The virus-infected white weed plants exhibit foliar vein swelling and yellow vein mosaic symptoms. Transmission The transmission of AYVSLV has not been investigated. It is likely that, in common with other begomoviruses, AYVSLV will be transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Similarly, it is likely that the virus will prove not to be either mechanically or seedtransmitted. Virion properties and genome The structure of the virions of AYVSLV has not been investigated. In common with all geminiviruses, the virions of AYVSLV are likely geminate (twinned icosahedra). With only a single isolate of AYVSLV having been characterized, the precise nature of the genome, whether monopartite, monopartite betasatellite-associated, or bipartite, is unclear. The characterized genome/genomic component of AYVSLV is 2748 nt (AF314144 = NC_002981) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). The Ageratum yellow vein Sri Lanka betasatellite DNA molecule is associated with AYVSLV, and consists of 1351 nt (AJ542498) (Bull et al. 2004; Zhou 2013).

Ageratum yellow vein virus Taxonomic position Genus: Begomovirus

(AYVV)

Family: Geminiviridae

Ageratum spp. (White weed)

41

Geographical distribution AYVV was first reported in Ageratum conyzoides plants from Singapore by Wong et al. (1993). Since then, the virus has also been identified in China, Nepal, the Philippines, Japan, Indonesia, Taiwan, Pakistan, India, Singapore, Vietnam, and Thailand (Swanson et al. 1993; Tan and Wong 1993; Tan et al. 1995; Stanley et al. 1997; Huang and Zhou 2006; Kon et al. 2007; Xiong et al. 2007; Jiao et al. 2013). Some isolates of the virus have previously been called Soybean crinkle leaf virus (Samretwanich et al. 2001), Ageratum yellow vein Hualian virus, and Ageratum yellow vein China virus (Xiong et al. 2007). Symptoms and host(s) The virus-infected white weed plants exhibit a bright foliar vein yellowing. However, care needs to be taken in interpreting symptoms since these are usually determined by the betasatellite with which the virus associates (Saunders et al. 2000). The virus has been identified in A. conyzoides, soybean (Glycine max), and tomato (Solanum lycopersicum). Transmission The virus was shown to be transmitted by the whitefly Bemisia tabaci between A. conyzoides plants. The virus is not transmitted by mechanical inoculation, not transmissible by contact between plants, and not transmissible by seed or pollen (Tan and Wong 1993; Tan et al. 1995). It is likely that, in common with other begomoviruses, AYVV is transmitted by B. tabaci in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 20  30 nm with a single coat protein (Tan and Wong 1993). The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2747 nt (AJ495813 = NC_004090; X74516 = NC_004626) (Tan et al. 1995; Stanley et al. 1997; Saunders and Stanley 1999; Briddon 2001; Samretwanich et al. 2001; Xiong et al. 2007; Brown et al. 2015; Zerbini et al. 2017). The genome of AYVV encodes the six genes typically encoded by monopartite begomoviruses and the DNA A component of bipartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated. The Ageratum yellow vein betasatellite DNA molecule is associated with AYVV, and consists of 1347 nt (AJ252072) (Saunders et al. 2000, 2004; Zhou 2013).

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

ArMV infection in plants of Ageratum spp. was reported from Iran (Ghotbi and Shahraeen 2005). The virus-infected white weed plants exhibit mosaic, leaf chlorosis, small necrotic lesions, leaf malformation, and deformation symptoms. This virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

A

42

Bidens mottle virus Taxonomic position Genus: Potyvirus

Ageratum spp. (White weed)

(BiMoV)

Family: Potyviridae

BiMoV infection in plants of Ageratum conyzoides was reported from Florida (Logan et al. 1984). The virus-infected ageratum plants showed stunting, leaf distortion, and flower deformation. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BiMoV, refer to Bidens spp.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Ageratum houstonianum was reported from Italy (Roggero et al. 1999). The virus-infected white weed plants exhibit symptoms of small necrotic rings and leaf malformation. The virus is transmitted by a thrips vector, Frankliniella occidentalis, in a persistent propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Malvastrum yellow vein virus Taxonomic position Genus: Begomovirus

(MaYVV)

Family: Geminiviridae

MaYVV infection in plants of Ageratum conyzoides was reported from China (Jiang and Zhou 2004). The virus-infected white weed plants exhibit yellow vein symptoms. No transmission data is available. However, the virus is expected to be transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The viral genomes the DNA-A of 2739 nt (AJ744881) and betasatellite of 1337 nt (AJ744882) from A. conyzoides were determined (Jiang and Zhou 2004). For more details of MaYVV, refer to Malvastrum spp.

Mungbean yellow mosaic India virus Taxonomic position Genus: Begomovirus

(MYMIV)

Family: Geminiviridae

MYMIV infection in plants of A. conyzoides was reported from India (Naimuddin et al. 2014). The virus-infected white weed plants exhibit yellow vein symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The virus is not transmitted by mechanical inoculation. For more details of MYMIV, refer to Vigna radiata.

Ageratum spp. (White weed)

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Pepper yellow leaf curl Indonesia virus Taxonomic position Genus: Begomovirus

(PepYLCIV)

A

Family: Geminiviridae

PepYLCIV infection in plants of Ageratum conyzoides was reported from Indonesia (Shibuya et al. 2007; Sakata et al. 2008). The virus-infected white weed plants exhibit yellow vein symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of PepYLCIV, refer to Capsicum annuum.

Sida yellow mosaic China virus Taxonomic position Genus: Begomovirus

(SiYMCNV)

Family: Geminiviridae

SiYMCNV infection in plants of Ageratum conyzoides was reported from China (Xiong and Zhou 2006). The virus-infected white weed plants exhibit yellow vein symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of SiYMCNV, refer to Sida spp.

Tomato leaf curl Ghana virus

(ToLCGV)

Synonyms Ageratum leaf curl Cameroon virus (ALCCMV) Taxonomic position Genus: Begomovirus

Family: Geminiviridae

ALCCMV infection in plants of Ageratum conyzoides was reported from Cameroon (Leke et al. 2012). The virus-infected white weed plants exhibit leaf curl symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Ageratum leaf curl Cameroon betasatellite DNA molecule is associated with ALCCMV, and consists of 1389 nt (NC_012557) (Zhou 2013). For more details of ToLCGV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Ageratum houstonianum was reported from California (Gumpf and Weathers 1972). The virus-infected white weed plants exhibit symptoms of mosaic patterns on severely distorted leaves and decline in vigor. The virus is transmitted by thrips vectors Frankliniella occidentalis and Thrips tabaci in a persistent propagative manner, and also by mechanical sapinoculation to a large number of vegetable and ornamental crops. For more details of TSWV, refer to Solanum lycopersicum.

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Ageratum spp. (White weed)

References Briddon RW (2001) Begomovirus. Geminiviridae The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Bull SE, Tsai WS, Briddon RW, Markham PG, Stanley J, Green SK (2004) Diversity of Begomovirus DNA beta satellites of non-malvaceous plants in east and south East Asia. Arch Virol 149(6):1193–1200 Ghotbi T, Shahraeen N (2005) First report on incidence of Arabis mosaic virus (ArMV, Nepovirus) on ornamental plants in Iran. Iranian J Plant Pathol 41(2):305–306 Gumpf DJ, Weathers LG (1972) Identification and purification of Tomato spotted wilt virus isolated from Ageratum. Plant Dis Reptr 56:859–872 Huang JF, Zhou XP (2006) Molecular characterization of two distinct begomoviruses from Ageratum conyzoides and Malvastrum coromandelianum in China. J Phytopathol 154:648–653 Jiang T, Zhou XP (2004) First report of Malvastrum yellow vein virus infecting Ageratum conyzoides. Plant Pathol 53:799 Jiao X, Gong H, Liu X, Xie Y, Zhou X (2013) Etiology of Ageratum yellow vein diseases in South China. Plant Dis 97:1497–1503 Khan MS, Tiwari AK, Ji SH, Chun SC (2012) Ageratum conyzoides and its role in begomoviral epidemics; Ageratum enation virus: an emerging threat in India. Vegetos 25(2):20–28 Khan MS, Tiwari AK, Raj SK, Srivastava A, Hye Ji S, Chun SC (2014) Molecular epidemiology of begomoviruses occurring on some vegetables, grain legume and weed species in the Terai belt of North India. Journal of Plant Diseases and Protection 121:53–57 Kon T, Kuwabara K, Hidayat SH, Ikegami M (2007) A Begomovirus associated with ageratum yellow vein disease in Indonesia: evidence for natural recombination between Tomato leaf curl Java virus and Ageratum yellow vein virus-[Java]. Arch Virol 152(6):1147–1157 Kumar Y, Hallan V, Zaidi AA (2011) First report of Ageratum enation virus infecting Crassocephalum crepidioides (Benth) S. Moore and Ageratum conyzoides L. in India. J Gen Plant Pathol 77:214–216 Kumar J, Gunapati S, Singh SP, Gadre R, Sharma NC, Tuli R (2013) Molecular characterization and pathogenicity of a carrot (Daucus carota) infecting begomovirus and associated betasatellite from India. Virus Res 178:478–485 Leke WN, Brown JK, Ligthart ME, Sattar N, Njualem DK, Kvarnheden A (2012) Ageratum conyzoides: a host to a unique begomovirus disease complex in Cameroon. Virus Res 163(1):229–237 Logan AE, Zettler FW, Christie SR (1984) Susceptibility of Rudbeckia, Zinnia, Ageratum and other bedding plants to Bidens mottle virus. Plant Dis 68:260–262 Naimuddin AM, Gupta S, Agnihotri AK (2014) Ageratum conyzoides Harbours Mungbean yellow mosaic India virus. Plant Pathol J 13:59–64 Pandey N, Tiwari AK, Rao GP, Shukla K (2011) Detection and identification of Ageratum enation virus infecting Ageratum conyzoides in India. Acta Phytopathologica et Entomologica Hungrica 46:203–212 Roggero P, Ciuffo M, Dellavalle G, Gotta P, Gallo S, Peters D (1999) Additional ornamental species as hosts of Impatiens necrotic spot tospovirus in Italy. Plant Dis 83:967 Sakata JJ, Shibuya Y, Sharma P, Ikegami M (2008) Strains of a new bipartite begomovirus, Pepper yellow leaf curl Indonesia virus, in leaf-curl-diseased tomato and yellow-vein-diseased ageratum in Indonesia. Arch Virol 153:2307–2313 Samretwanich K, Kittipakorn K, Chiemsombat P, Ikegami M (2001) Complete nucleotide sequence and genome organization of soybean crinkle leaf virus. J Phytopathol 149:333–336 Saunders K, Stanley J (1999) A nanovirus-like component associated with yellow vein disease of Ageratum conyzoides: evidence for inter-family recombination between plant DNA viruses. Virology 264:142–152 Saunders K, Bedford ID, Briddon RW, Markham PG, Wong SM, Stanley J (2000) A unique virus complex causes Ageratum yellow vein disease. Proc Natl Acad Sci USA 97:6890–6895 Saunders K, Norman A, Gucciardo S, Stanley J (2004) The DNA b satellite component associated with ageratum yellow vein disease encodes an essential pathogenicity protein (bC1). Virology 324:37–47 Scott SW (2011) Ilarvirus. Bromoviridae. The Springer index of viruses. Springer, New York, pp 187–194. https://doi. org/10.1007/978-0-387-95919-1_27 Sharman M, Thomas JE (2013) Genetic diversity of subgroup 1 Ilarviruses from eastern Australia. Arch Virol 158:1637–1647 Shibuya Y, Sakata J, Sukamto N, Kon T, Sharma P, Ikegami M (2007) First report of Pepper yellow leaf curl Indonesia virus in Ageratum conyzoides in Indonesia. Plant Dis 91:1198 Srivastava A, Raj SK, Kumar S, Snehi SK, Kulshreshtha A, Hallan V, Pande SS (2013) Molecular identification of Ageratum enation virus, betasatellite and alphasatellite molecules isolated from yellow vein diseased Amaranthus cruentus in India. Virus Genes 47(3):584–590

Aglaonema spp.

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Srivastava A, Kumar S, Raj SK (2015) Molecular characterization of a Begomovirus and betasatellite causing yellow vein net disease of Ageratum houstonianum. Plant Dis 99(5):627–631 Stanley J, Saunders K, Pinner MS, Wong SM (1997) Novel defective interfering DNAs associated with Ageratum yellow vein geminivirus infection of Ageratum conyzoides. Virology 239:87–96 Swanson MM, Harrison BD, Wong SM (1993) A geminivirus causing vein yellowing of Ageratum conyzoides in Singapore. Plant Pathol 42:137–139 Tahir M, Amin I, Haider MS, Mansoor S, Briddon RW (2015) Ageratum enation virus – a begomovirus of weeds with the potential to infect crops. Viruses 7:647–665 Tan HNP, Wong SM (1993) Some properties of Singapore Ageratum yellow vein virus (SAYVV). J Phytopathol 139:165–176 Tan PHN, Wong SM, Wu M, Bedford ID, Saunders K, Stanley J (1995) Genome organization of Ageratum yellow vein virus, a monopartite whitefly-transmitted geminivirus isolated from a common weed. J Gen Virol 76:2915–2922 Tsai WS, Shih SL, Kenyon L, Green SK, Jan F-J (2011) Temporal distribution and pathogenicity of the predominant tomato-infecting begomoviruses in Taiwan. Plant Pathol 60:787–799 Wong SM, Swanson MM, Harrison BD (1993) A new geminivirus causing vein yellowing of Ageratum conyzoides in Singapore. Plant Pathol 42B:137–139 Xiong Q, Zhou XP (2006) First report of Sida yellow mosaic China virus associated with yellow vein disease of Ageratum conyzoides in China. J Plant Pathol 88:125–125 Xiong Q, Fan S, Wu J, Zhou X (2007) Ageratum yellow vein China virus is a distinct begomovirus species associated with a DNAb molecule. Phytopathology 97:405–411 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zhou X (2013) Advances in understanding begomovirus satellites. Annu Rev Phytopathol 51:357–381

Aglaonema spp. Family: Araceae

Ornamental

Aglaonema bacilliform virus Taxonomic position Genus: Badnavirus

(ABV)

Family: Caulimoviridae

Geographical distribution ABV infection in plants of Aglaonema modestum was reported from Minnesota, USA (Lockhart 1996). Symptoms and host(s) The virus-infected Aglaonema plants exhibit symptoms of leaf mottling, chlorosis, deformation, green ring-spots, and premature senescence. Transmission The virus is not transmitted by the citrus mealybug Planococcus citri. Virion properties and genome The virions are bacilliform in shape with a diameter of 30 nm and modal particle length of 130 nm. The genome is a monopartite, circular double-stranded DNA of 7000–7600 bp with a single-strand discontinuity at one site in each strand (Olszewski and Lockhart 2011; Bhat et al. 2016).

A

46

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Aglaonema spp.

(AMV)

Family: Bromoviridae

AMV infection in plants of Aglaonema spp. was reported from New Zealand (Fletcher 1983). The virus-infected Aglaonema plants exhibit symptoms of interveinal yellow flecking on leaves. The virus is transmitted by a large number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Dasheen mosaic virus Taxonomic position Genus: Potyvirus

(DsMV)

Family: Potyviridae

DsMV infection in plants of Aglaonema spp. was reported from India and the USA (Elliott et al. 1997; Ram et al. 2003). The virus-infected Aglaonema plants exhibit systemic mosaic symptoms (Nelson 2008). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of DsMV, refer to Colocasia esculenta.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Aglaonema commutatum was reported from Pennsylvania (USA) (Hausbeck et al. 1992). The virus-infected Aglaonema plants exhibit necrosis and chlorotic spotting symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting, but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Bhat AI, Hohn T, Selvarajan R (2016) Badnaviruses: the current global scenario. Viruses 8(6):177 Elliott MS, Zettler FW, Brown LG (1997) Dasheen mosaic potyvirus of edible and ornamental aroids. Plant pathology circular no. 384. Florida Department of Agriculture and Consumer Services, Florida, July/Aug 1997 Fletcher JD (1983) New plant disease records in New Zealand: additional hosts of Alfalfa mosaic virus. N Z J Agric Res 28:279–282 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among green house ornamentals in Pennsylvania. Plant Dis 76:795–800 Lockhart BEL (1996) Association of a Badnavirus with premature leaf senescence in Aglaonema. Phytoparasitica 24:33–334 Nelson SC (2008) Dasheen mosaic of edible and ornamental aroids. Plant Disease CTAHR. Available at: http://www. ctahr.hawaii.edu/oc/freepubs/pdf/PD-44.pdf Olszewski NE, Lockhart B (2011) Badnavirus. Caulimoviridae. In: The Springer index of viruses. Springer, New York, pp 263–269. https://doi.org/10.1007/978-0-387-95919-1_40 Ram R, Joshi A, Verma N, Kulshrestha S, Raikhy G, Hallan V, Zaidi AA (2003) First report of Dasheen mosaic virus infecting four ornamental aroids in India. Plant Pathol 52:411

Agropyron repens (Couch grass)

47

Agropyron repens (Couch grass) A

Synonyms Elymus repens Family: Poaceae

Forage crop

Agropyron mosaic virus Taxonomic position Genus: Rymovirus

(AgMV)

Family: Potyviridae

Geographical distribution AgMV was first reported in plants of Agropyron repens from the USA (McKinney 1937). The virus spreads in the USA, southern Canada, Finland, Germany, Great Britain, and Hungary (Bremer 1964; Shepard 1968; Catherall and Chamberlain 1975; Seifers 1992). Symptoms and host(s) The two main symptom types of streak mosaic that occur in AgMV-infected A. repens leaves are palegreen streaks that are masked during plant maturation and yellow chlorotic streaks that become highly pronounced on older leaves, and in general infected plants are stunted. The natural hosts of this virus are Aristida spp., Bothriochloa laguroides, Elymus smithii, Elytrigia repens, E. intermedia, Lophopyrum elongatum, Secale cereale, Sorghastrum nutans, Triticum aestivum, and T. durum. Transmission The virus is transmitted by the mite vector, Abacarus hystrix in a semi-persistent manner. Not transmitted by the mite species Aculus mckenziei and Eriophyes tulipae. The virus is transmissible by mechanical sap-inoculation to 17 other festucoid species of Gramineae, not transmitted by seed, and is not transmitted by pollen (Slykhuis 1969). Virion properties and genome The virions are non-enveloped, flexuous filaments, 690–720 nm long, and 15 nm in diameter. The virions contain a single molecule of linear positive-sense ssRNA of 9540 nt (AY623626 = NC_005903) with a 30 -poly(A) terminus (Seifers 1992; Zagula et al. 1992; Salm et al. 1996; French and Stenger 2005; LopezMoya et al. 2009; Wylie et al. 2017).

References Bremer K (1964) Agropyron mosaic virus in Finland. Ann Agric Fenn 3:324–333 Catherall PL, Chamberlain JA (1975) Occurrence of Agropyron mosaic virus in Britain. Plant Pathol 24:155–157 French R, Stenger DC (2005) Genome sequences of Agropyron mosaic virus and Hordeum mosaic virus support reciprocal monophyly of the genera Potyvirus and Rymovirus in the family Potyviridae. Arch Virol 150:299–312

48

Ajuga reptans and A. genevensis (Upright bugle)

Lopez-Moya JJ, Valli A, Garcia JA (2009) Potyviridae. In: Iacobellis NS, Collmer A, Hutcheson SW, Mansfield JW, Morris CE, Murillo J, Schaad D NW, Stead E, Surico G, Ullrich M (eds.). Encyclopaedia of life sciences (ELS). Kluwer Academic, Dordrecht, The Netherlands. https://doi.org/10.1002/9780470015902.a0000755.pub2 McKinney HH (1937) Mosaic disease of wheat and related cereals. US Dept Agr Circular No. 442, p 23 Salm SN, Rey MEC, Robertson NL, French R, Rabenstein F, Schubert J (1996) Molecular cloning and nucleotide sequencing of the partial genomes of Agropyron and Hordeum mosaic viruses, two members of the Rymovirus genus in the taxonomic family Potyviridae. Arch Virol 141:2115–2127 Seifers DL (1992) Partial characterization of a Colorado isolate of Agropyron mosaic virus. Plant Dis 76:564–569 Shepard JF (1968) Occurrence of Agropyron mosaic virus in Montana. Plant Dis Reptr 52:139–141 Slykhuis JT (1969) Transmission of Agropyron mosaic virus by the eriophyid mite, Abacarus hystrix. Phytopathology 59:29–32 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Zagula KR, Niblett CL, Robertson NL, French R, Lommen SA (1992) Potyviridae: genus Rymovirus. Arch Virol 5:269–276

Ajuga reptans and A. genevensis (Upright bugle) Family: Lamiaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Medicinal

(AMV)

Family: Bromoviridae

AMV infection in plants of Ajuga reptans was reported from Australia and the USA (Schroeder and Provvidenti 1972; Shukla and Gough 1983; Fisher and Nameth 2000). The virus-infected upright bugle plants exhibit mild ring-spot and mottle symptoms. The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Broad bean wilt virus Taxonomic position Genus: Fabavirus

(BBWV)

Family: Secoviridae

BBWV (but without specifying either of the currently recognized species BBWV-1 or BBWV-2) infection in plants of Ajuga reptans was reported from Australia (Shukla and Gough 1983). The virus-infected upright bugle plants exhibit symptoms of chlorotic and necrotic flecking. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BBWV, refer to Vicia faba.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

Ajuga reptans and A. genevensis (Upright bugle)

49

CMV infection in plants of Ajuga reptans was reported from Australia and Ohio (USA) (Shukla and Gough 1983; Fisher and Nameth 1997, 2000, 2003). The virus-infected upright bugle plants exhibit mosaic symptoms on leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Plum pox virus

(PPV)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

PPV infection in plants of Ajuga genevensis was reported from Slovenia (Virscek Marn et al. 2004). The virus-infected upright bugle plants exhibit pale or yellow-green ring-spots or mottling on the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of PPV, refer to Prunus domestica.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Ajuga reptans was reported from Ohio (Fisher 2013). The virus-infected upright bugle plants exhibit subtle virus-like mottle symptoms. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Ajuga reptans was reported from Australia and Ohio (USA) (Shukla and Gough 1983; Fisher and Nameth 1997, 2000). The virus-infected upright bugle plants exhibit symptoms of mottling and crinkling of the leaves. The virus is transmitted by thrips vectors, the virus present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible. The virus is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

References Fisher JR (2013) First report of Tobacco ringspot virus infecting Ajuga reptans in Ohio. Online. Plant Health Prog. https:// doi.org/10.1094/PHP-2013-0301-01-BR Fisher JR, Nameth ST (1997) Cucumber mosaic virus, Tobacco streak virus, and Cucumber mosaic virus satellite RNA associated with mosaic and ringspot symptoms in Ajuga reptans in Ohio. Plant Dis 81:1214 Fisher JR, Nameth ST (2000) Virus assessment of Ajuga reptans cultivars reveals Alfalfa mosaic, Tobacco streak, and Cucumber mosaic (CMV) viruses, and a CMV satellite RNA. Hortic Sci 35:230–234

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50

Allamanda cathartica (Allamanda)

Fisher JR, Nameth SGP (2003) Characterization of a Cucumber mosaic virus isolate and satellite RNA from the ornamental host Ajuga reptans ‘Royalty’. J Amer Soc Hortic Sci 128(2):231–237 Schroeder WT, Provvidenti R (1972) Alfalfa mosaic virus in Ajuga reptans. Plant Dis Rptr 56:285 Shukla DD, Gough KH (1983) Tobacco streak, broad bean wilt, cucumber mosaic and alfalfa mosaic viruses associated with ring spot of Ajuga reptans in Australia. Plant Dis 67:221–224 Virscek Marn M, Mavric I, Urbancic-Zemljic M, Skerlavaj V (2004) Detection of Plum pox virus potyvirus in weeds. Acta Hortic 657:251–254

Allamanda cathartica (Allamanda) Family: Fabaceae

Ornamental

Allamanda leaf curl virus Taxonomic position Genus: Begomovirus

(AllLCV)

Family: Geminiviridae

Geographical distribution AllLCV infection in plants of Allamanda cathartica was reported from China (He et al. 2009). Symptoms and host(s) The virus-infected allamanda plants exhibit leaf curling and swelling of veins. Transmission The transmission of AllLCV has not been investigated. It is likely that, in common with other begomoviruses, AllLCV will be transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Similarly, it is likely that the virus will prove not to be either mechanically or seed-transmitted. Virion properties and genome The structure of the virions of AllLCV has not been investigated. In common with all geminiviruses, the virions of AllLCV are likely geminate (twinned icosahedra). With only a single isolate of AllLCV having been characterized, the precise nature of the genome (whether monopartite, monopartite betasatellite-associated, or bipartite) is unclear. The characterized genome/genomic component of AllLCV is 2755 nt (EF602306 = NC_010947). The characterized genome of AllLCV encodes the six genes typically encoded by monopartite begomoviruses. The expression and function of the genes have not been investigated. The virus does not appear to associate with a betasatellite (Briddon 2001; He et al. 2009; Brown et al. 2015; Zerbini et al. 2017).

Allamanda leaf mottle distortion virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

(AllLMoDV)

Allamanda cathartica (Allamanda)

51

Geographical distribution AllLMoDV infection in plants of Allamanda cathartica was reported from India (Shilpi et al. unpublished – KC202818; Jailani et al. unpublished – MG969497). Symptoms and host(s) The virus-infected golden trumpet plant leaves show mottling and distortion symptoms. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm, with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2772 nt (KC202818 = NC_024009), and DNA-B of 2690 nt (MG969497) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Allamanda cathartica was reported from Brazil, Italy, and Taiwan (Rodrigues et al. 1984; Bellardi and Bertaccini 1993; Alexandre et al. 2005; Chen et al. 2005; Chen and Yang 2006). The virus-infected allamanda plants exhibit symptoms of mottle and yellow spots or severe mosaic accompanied by rugosity and leaf distortion. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

References Alexandre MAV, Rivas EB, Tozetto ARP, Duarte LML (2005) An annotated list on the natural occurrence of viruses in ornamental plants in Brazil. Instituto Biologico, Sao Paulo, p 54 Bellardi MG, Bertaccini A (1993) First report of Cucumber mosaic virus (CMV) in Allamanda plants and in Delphinium seed. In: Abstracts, 6th international congress of plant pathology, July 28–August 6, 1993, Montreal, No. 17.1.2, 301 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Chen YK, Yang CC (2006) Characterization of an isolate of Cucumber mosaic virus associated with allamanda mosaic. Plant Pathol Bull 15:77–82 Chen YK, Yang CC, Hsu HT (2005) Allamanda mosaic caused by Cucumber mosaic virus in Taiwan. Plant Dis 89(5):529 He ZF, Mao MJ, Yu H, Li HP (2009) Molecular characterization of a distinct begomovirus infecting Allamanda cathartica in Guangdong, China. Arch Virol 154:1199–1202 Rodrigues MGR, Kitajima EW, Lin MT (1984) Ringspot of Allamanda cathartica associated with Cucumber mosaic virus infection. Fitopatol Bras 9(1):151–154 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

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Alliaria spp. (Garlic mustard)

Alliaria spp. (Garlic mustard) Family: Brassicaceae

Leafy vegetable

Cucumber mosaic virus

(CMV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV infection in plants of Alliaria officinalis was reported from Ohio, USA (Boehm and Nameth 2000). The virus-infected garlic mustard plants exhibit leaf mosaic and vein banding symptoms. The virus is transmitted by several aphid species in a non-persistent manner, and also by mechanical sapinoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Alliaria petiolata was reported from Southern Ontario, Minnesota, Iran, and Europe (Lisa and Lovisolo 1976; Stobbs and Van Schagen 1987; Lockhart 2012; Zhao et al. 2016; Farzadfar and Pourrahim 2017). The virus-infected garlic mustard plants exhibit severe mottling with dark green areas, leaf puckering, marginal downward leafroll, some veinal necrosis, and general stunting of the plant. Symptoms were more pronounced in plants that received diffuse sun than those growing in full shade. The virus is transmitted by an aphid vector Myzus persicae in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TuMV, refer to Brassica rapa.

White clover mosaic virus Taxonomic position Genus: Potexvirus

(WClMV)

Family: Alphaflexiviridae

WClMV infection in plants of Alliaria petiolata was reported from Pennsylvania (Zhao et al. 2016). The virus-infected garlic mustard plants exhibit mosaic symptoms. The virus is sap-transmissible and also through contact between plants. For more details of WClMV, refer to Trifolium spp.

References Boehm MJ, Nameth ST (2000) First report of Cucumber mosaic virus in garlic mustard in Ohio. Plant Dis 84:1047 Farzadfar S, Pourrahim R (2017) First report of Turnip mosaic virus infection of Alliaria petiolata in Iran. Plant Dis 101:1558 Lisa V, Lovisolo O (1976) Biological and serological characterization of the Alliaria strain of Turnip mosaic virus. Phytopathol Z 86:90–96

Allium ampeloprasum var. porrum (Leek)

53

Lockhart BEL (2012) First report of Turnip mosaic virus occurrence in garlic mustard in Minnesota. Online. Plant Health Prog. https://doi.org/10.1094/PHP-2012-08XX-01-BR Stobbs LW, Van Schagen JG (1987) Occurrence and characterization of a Turnip mosaic virus isolate infecting Alliaria petiolata in Ontario, Canada. Plant Dis 71:965–968 Zhao K, Margaria P, Rosa C (2016) First report of White clover mosaic virus and Turnip mosaic virus mixed infection in garlic mustard in Pennsylvania. Plant Dis 100:866

Allium ampeloprasum var. porrum (Leek) Family: Amaryllidaceae

Spice crop

Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

IYSV was reported on plants of Allium ampeloprasum var. porrum in the Netherlands, Spain, Greece, Germany, Holland, Austria, Brazil, India, Australia, Japan, South Africa, Peru, Mexico, Chile, Slovenia, Reunion Island, Spain, Zimbabwe, and the USA (Anon 1998; Cordoba-Selles et al. 2007; Gent et al. 2007; Schwartz et al. 2007; Chatzivassiliou et al. 2009; Gamage et al. 2010; Karavina et al. 2016; Karavina and Gubba 2017). On leek IYSV produces symptoms of elongated, oval chlorotic rings that turn yellow and eventually become necrotic. Rings could occasionally be observed overlapping each other. In some cases, symptoms were said to be chlorotic spots that later became necrotic (Mavric and Ravnikar 2001). The virus is transmitted by onion thrips, Thrips tabaci, in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of IYSV, refer to Iris spp. or Allium cepa.

Leek white stripe virus Taxonomic position Genus: Betanecrovirus

(LWSV)

Family: Tombusviridae

Geographical distribution LWSV infection in plants of Allium ampeloprasum var. porrum was reported from France (Lot et al. 1996). Symptoms and host(s) The virus-infected leek plants exhibit white stripes on the leaves that extend to the stem; occasionally some plants turn completely white and rot (Lot et al. 1996). Transmission No vector is identified for this virus. The virus is mechanically sap-transmissible with difficulty and has a restricted host range. The virus is possibly soil-borne.

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54

Allium ampeloprasum var. porrum (Leek)

Virion properties and genome The virions are isometric 28 nm in diameter and have T = 3 icosahedral symmetry. The genome is a monopartite, linear, single-stranded positive-sense RNA of 3662 nt (X94560 = NC_001822) (Lot et al. 1996).

Leek yellow stripe virus

(LYSV)

Synonyms Garlic mosaic virus; Pterostylis virus Y Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution LYSV was first reported from Germany and European countries infecting Allium porrum (Bos et al. 1978). This virus occurs in the Eurasian region, South America, Turkey, New Zealand, Iran, Serbia, and Australia (Sward 1991; Hall et al. 2007; Shahraeen et al. 2008; Fidan and Baloglu 2009; Korkmaz and Cevik 2009; Vucurovic et al. 2016). Symptoms and host(s) The virus-infected leek plants exhibit symptoms of necrotic or chlorotic streaks and a general gray lusterless color. Even mosaic, yellow stripe, malformation, dwarfing, and wilting symptoms are also recorded (Paludan 1980; Graichen 1991). Transmission The virus is transmitted by aphid vectors, Myzus persicae, Aphis fabae, Aphis gossypii, Aphis nerii, Hyperomyzus carduellinus, Rhopalosiphum maidis, R. padi, Schizaphis graminum, and Uroleucon sonchi in a non-persistent manner (Lunello et al. 2002). The virus is transmissible by mechanical sapinoculation to Allium spp. and Chenopodium spp.; Chenopodium amaranticolor, C. quinoa, C. murale, and C. album produce chlorotic and necrotic local lesions on the inoculated leaves. Onion is rarely infected, and shallot is immune to this virus (Lunello et al. 2002). The virus is not transmissible by contact between plants and not transmitted by seed. Virion properties and genome The virions are non-enveloped and flexuous filaments, with a clear modal length of 800–820 nm by 13 nm. The genome consists of a single molecule of linear positive-sense single-stranded RNA of 10142 nt (AJ307057 = NC_004011) (Chen et al. 2002; Takaki et al. 2005; Revers and Garcia 2015; Wylie et al. 2017).

Onion yellow dwarf virus Taxonomic position Genus: Potyvirus

(OYDV)

Family: Potyviridae

OYDV infection in plants of Allium ampeloprasum var. porrum was reported from Australia, Spain, and Iran (Hall et al. 2007; Shahraeen et al. 2008; Naderi Saffar et al. 2013; Fernandez-Tabanera et al. 2018).

Allium ampeloprasum var. porrum (Leek)

55

The virus-infected leek plants exhibit symptoms of mosaic, yellow streak striping, curling, and distortion of flower stems. The virus is transmitted by a number of aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of OYDV, refer to Allium cepa.

Shallot virus X

(ShVX)

Synonyms Shallot mite-borne latent virus (SMbLV)

Taxonomic position Genus: Allexivirus

Family: Alphaflexiviridae

ShVX infection in plants of Allium ampeloprasum var. porrum was reported from New Zealand (Van Dijk and Van Der Vlugt 1994). The virus is transmitted by a mite vector, Aceria tulipae, and also by mechanical sap-inoculation. The use of infected propagative material is another source of virus spread. For more details of ShVX, refer to Allium cepa var. aggregatum.

Tomato black ring virus Taxonomic position Genus: Nepovirus

(TBRV)

Family: Secoviridae

TBRV infection in plants of Allium ampeloprasum var. porrum was reported from Northern Ireland (Calvert and Harrison 1963). The virus-infected leek plants exhibit symptoms of irregular longitudinal chlorotic markings and stunting. The virus is transmitted by the nematode vector Longidorus attenuatus in a non-persistent manner, and also by mechanical sap-inoculation (Brown and Trudgill 1998). For more details of TBRV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Allium ampeloprasum var. porrum was widespread in the Mediterranean basin and the USA (Nischwitz et al. 2006). The virus-infected leek plants exhibited extended bleaching of leaf tips and necrotic lesions. The virus is transmitted by thrips vectors, Frankliniella schulzei, Frankliniella occidentalis, and Thrips tabaci, in a persistent-propagative manner (Chatzivassiliou et al. 1999). The virus is mechanically sap-transmissible to a number of hosts. The virus is transmissible by grafting, but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

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Turnip mosaic virus Taxonomic position Genus: Potyvirus

Allium ampeloprasum var. porrum (Leek)

(TuMV)

Family: Potyviridae

TuMV infection in plants of Allium ampeloprasum var. porrum was reported from Israel, Yugloslavia, and Greece (Stefanac and Plese 1980; Gera et al. 1997; Dovas et al. 2001). The virus-infected leek plants show reduction of plant growth and yellow striping in the leaves. The virus is transmitted by aphid vectors, Myzus persicae and Aphis gossypii, in a non-persistent manner, and also by mechanical sap-inoculation to a large number of test plants. For more details of TuMV, refer to Brassica rapa.

References Anon (1998) Iris yellow spot virus in leek. In: Annual report 1997, Diagnostic Centre. Plant Protection Service, the Netherlands. pp 118–119 Bos L, Huijberts N, Huttingaand H, Maat DZ (1978) Leek yellow stripe virus and its relationships to Onion yellow dwarf virus: characterization, ecology and possible control. Neth J Plant Pathol 84:185–204 Brown DJF, Trudgill DL (1998) Nematode transmission of plant viruses: a 30 year perspective. Host pathogen interactions and crop protection. SCRI Annual Report, pp 121–125 Calvert EL, Harrison BD (1963) Outbreaks of Tomato black ring virus in onion and leek in Northern Ireland. Hortic Res 2:115–120 Chatzivassiliou EK, Nagata T, Katis NI, Peters D (1999) Transmission of Tomato spotted wilt tospovirus by Thrips tabaci populations originating from leek. Plant Pathol 48:700–706 Chatzivassiliou EK, Giavachtsia V, Mehraban AH, Hoedjes K, Peters D (2009) Identification and incidence of Iris yellow spot virus, a new pathogen in onion and leek in Greece. Plant Dis 93:761 Chen J, Chen JP, Adams MJ (2002) Characterisation of some carla- and potyviruses from bulb crops in China. Brief report. Arch Virol 147(2):419–428 Cordoba-Selles C, Cebrian-Mico C, Alfaro-Fernadez A (2007) First report of Iris yellow spot virus in commercial leek (Allium porrum) in Spain. Plant Dis 91:1365 Dovas CI, Hatziloukas E, Salomon R, Barg E, Shiboleth Y, Katis NI (2001) Incidence of viruses infecting Allium spp. in Greece. Eur J Plant Pathol 107:677–684 Fernandez-Tabanera E, Fraile A, Lunello P, Garcia-Arenal F, Ayllon MA (2018) First report of Onion yellow dwarf virus in Leek (Allium ampeloprasum var. porrum) in Spain. Plant Dis 102:256 Fidan H, Baloglu S (2009) First report of Leek yellow stripe virus on leek in Turkey. J Plant Pathol 91:234 Gamage SMKW, Hassani-Mehraban A, Peters D (2010) Identification of Iris yellow spot virus on leek (Allium porrum) in Sri Lanka. Plant Dis 94(8):1070 Gent DH, Martin RR, Ocamb CM (2007) First report of Iris yellow spot virus on onion and leek in western Oregon. Plant Dis 91:468 Gera A, Lesemann DE, Cohen J, Franck A, Levy S, Salomon R (1997) The natural occurrence of Turnip mosaic potyvirus in Allium ampeloprasum. J Phytopathol 145:289–293 Graichen K (1991) Gelbstreifigkeit des Porrees verursacht erhebliche Pflanzenausfälle. Gartenbauwissenschaft 38:17–19 Hall BH, Hitch CJ, Oxspring EA, Wicks TJ (2007) Leek diseases in Australia. Australas Plant Pathol 36:383–388 Karavina C, Gubba A (2017) Iris yellow spot virus in Zimbabwe: incidence, severity and characterization of Alliuminfecting isolates. Crop Prot 94:69–76 Karavina C, Ibaba JD, Gubba A, Pappu HR (2016) First report of Iris yellow spot virus infecting garlic and leek in Zimbabwe. Plant Dis 100:657 Korkmaz S, Cevik B (2009) Leek yellow stripe virus newly reported in Turkey. Plant Pathol 58:787 Lot H, Rubino L, Delecolle B, Jacquemond M, Turturo C, Russo M (1996) Characterization, nucleotide sequence and genome organization of Leek white stripe virus, a putative new species in the genus Necrovirus. Arch Virol 141:2375–2385 Lunello P, Ducasse DA, Helguera M, Nome SF, Conci VC (2002) An Argentinean isolate of Leek yellow stripe virus from leek can be transmitted to garlic. J Plant Pathol 84:11–17 Mavric I, Ravnikar M (2001) Iris yellow spot tospovirus in Slovenia. In: Catara A, Albanese G, Catara V, La Rosa R, Polizzi G, Tessitori M (eds) Proceedings of the 5th congress of the European Foundation for Plant Pathology: biodiversity in plant pathology, Taormina–Giardini Naxos, 18–22 Sept 2000. Societa Italiana di Patologia Vegetale, pp 223–225

Allium cepa (Onion)

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Naderi Saffar Z, Torabi S, Naghavi M, Golnaraghi AR, Aryakia E (2013) Onion yellow dwarf virus on leek, onion and welsh onion in Iran. J Plant Pathol 95:S4.73 Nischwitz C, Mullis SW, Gitaitis RD, Csinos AS (2006) First report of Tomato spotted wilt virus in Leek (Allium porrum) in the United States. Plant Dis 90:525 Paludan N (1980) Virus attack on leek: survey, diagnosis, tolerance of varieties and winter hardiness. Tidsskr Planteavl 84:371–385 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Schwartz HF, Otto K, Pappu HR (2007) First report of Iris yellow spot virus in Commercial Leek (Allium porrum) in the United States. Plant Dis 91:113 Shahraeen N, Lesemann DE, Ghotbi T (2008) Survey for viruses infecting onion, garlic and leek crops in Iran. OEPP/ EPPO Bull 38:131–135 Stefanac Z, Plese N (1980) Turnip mosaic virus in two Mediterranean Allium species. In: Proceedings of the fifth congress of the Mediterranean phytopathological union, Patras, 21–27 Sept, pp 37–38 Sward RJ (1991) Leek yellow stripe virus recorded in leek in Australia. Australas Plant Pathol 20:14–15 Takaki F, Sano T, Yamashita K, Fujita T, Ueda K, Kato T (2005) Complete nucleotide sequence of attenuated and severe isolate of Leek yellow stripe virus from garlic in Northern Japan: identification of three distinct virus types in garlic and leek worldwide. Arch Virol 150(16):1135–1149 Van Dijk P, Van Der Vlugt RAA (1994) New mite-borne virus isolates from rakkyo, shallot and leek species. Eur J Plant Pathol 100:269–277 Vucurovic I, Vucurovic A, Nikolic D, Bulajic A, Milosevic D, Krstic B, Stankovic I (2016) First report of Leek yellow stripe virus in Leek in Serbia. Plant Dis 100:230 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Allium cepa (Onion) Family: Amaryllidaceae

Spice crop

Allium cepa amalgavirus 1 Taxonomic position Genus: Amalgavirus

(AcAV1)

Family: Amalgaviridae

Geographical distribution AcAV1 infection in plants of Allium cepa was reported from the USA (Nibert et al. 2016). Transmission Transmission has not been proven, but AcAV1 appears likely to be transmitted efficiently through seed. Virion properties and genome No virion has been reported for amalgaviruses to date. The genome is double-stranded RNA of 3453 bp (BK010347 = NC_036580) and contains two overlapping ORFs (Nibert et al. 2016).

Allium cepa amalgavirus 2 Taxonomic position Genus: Amalgavirus

(AcAV2)

Family: Amalgaviridae

A

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Allium cepa (Onion)

Geographical distribution AcAV2 infection in plants of Allium cepa was reported from the USA (Nibert et al. 2016).

Transmission Transmission has not been proven, but appears likely to be transmitted efficiently through seed.

Virion properties and genome No virion has been reported for amalgaviruses to date. The genome is double-stranded RNA of 3453 bp (BK010348 = NC_036581) and contains two overlapping ORFs (Nibert et al. 2016).

Artichoke yellow ringspot virus Taxonomic position Genus: Nepovirus

(AYRSV)

Family: Secoviridae

AYRSV infection in plants of Allium cepa was reported from Greece (Maliogka et al. 2006). The virusinfected onion plants exhibit symptoms of yellow leaf striping and necrotic tips. No vector is known for this virus. The virus is mechanically sap-transmissible. The virus is seed-transmitted in onion (Maliogka et al. 2006). For more details of AYRSV, refer to Cynara cardunculus var. scolymus.

Garlic common latent virus Taxonomic position Genus: Carlavirus

(GarCLV)

Family: Betaflexiviridae

GarCLV infection in plants of Allium cepa was reported from Japan, Turkey, and Iran (Shahraeen et al. 2008; Sevik and Akcura 2013). The virus-infected onion plants do not exhibit any symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of GarCLV, refer to Allium sativum.

Garlic virus C Taxonomic position Genus: Allexivirus

(GarV-C)

Family: Alphaflexiviridae

GarV-C infection in plants of Allium cepa was reported from Iran (Shahraeen et al. 2008). The virusinfected onion plants exhibit growth reduction, twisting, and leaf malformation. The virus is transmitted by a mite vector, Aceria tulipae, and also by mechanical sap-inoculation. The use of infected vegetative propagative material is another source of virus spread. For more details of GarV-C, refer to Allium sativum.

Allium cepa (Onion)

Garlic virus D Taxonomic position Genus: Allexivirus

59

(GarV-D)

A

Family: Alphaflexiviridae

GarV-D infection in plants of Allium cepa was reported from Iran and India (Shahraeen et al. 2008; Khan et al. 2015). The virus-infected onion plants exhibit growth reduction, twisting, and leaf malformation. The virus is transmitted by a mite vector, Aceria tulipae, and also by mechanical sapinoculation. The use of infected vegetative propagative material is another source of virus spread. For more details of GarV-D, refer to Allium sativum.

Groundnut bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(GBNV)

Family: Tospoviridae

GBNV infection in plants of Allium cepa was reported from Kadapa district of Andhra Pradesh (India) (Sujitha et al. 2012). The virus-infected onion plants exhibit straw-colored mosaic and necrotic lesions on the young leaves. The necrosis starts with the apical portion of young leaves and flower stalks and finally causes flower distortion and plant death. The virus is transmitted by thrips vectors in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of GBNV, refer to Arachis hypogaea.

Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

Geographical distribution LYSV infection in plants of Allium cepa was reported from Brazil, Canada, France, Israel, Germany, Japan, Greece, Slovenia, Iran, Italy, Tunisia, Egypt, India, Pakistan, Peru, Chile, Uruguay, Guatemala, the USA, the UK, Africa, Spain, Austria, Australia, Serbia, New Zealand, Georgia, Zimbabwe, Mexico, and the Netherlands (Gera et al. 1998; Pozzer et al. 1999; Schwartz et al. 2002; Coutts et al. 2003; Creamer et al. 2004; Mullis et al. 2004, 2006; Ravi et al. 2006; Huchette et al. 2008; Bulajic et al. 2009; Ward et al. 2009b; Colnago et al. 2010; Lobin et al. 2010; Sether et al. 2010; Birithia et al. 2011; Kunkalikar et al. 2011; Plenk and Grausgruber-Groger 2011; Bag et al. 2012; Hoepting and Fuchs 2012; Kumar et al. 2012; Manglli et al. 2012; Rafizadeh et al. 2013; Iftikhar et al. 2013; Trkulja et al. 2013; Karavina et al. 2016; Velasquez-Valle et al. 2016; Karavina and Gubba 2017). Symptoms and host(s) The virus-infected onion plants exhibit characteristic symptoms of eyespot- to diamond-shaped lesions and lesions yellow- to straw-colored. Lesions will appear on both the seedstalk and the leaves. Seedstalks may swell at the point of infection. Late in the season, infected flower/seedstalks and leaves will lodge (fall over). Onions, garlic, leek, iris, lisianthus, weeds, jimsonweed, tobacco, and redroot pigweed are infected by this virus (Gent et al. 2006; Hsu et al. 2011).

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Allium cepa (Onion)

Transmission The virus is transmitted by onion thrips (Thrips tabaci) but is not transmitted by other common Thrips species, such as Western flower thrips (Frankliniella occidentalis) (Kritzman et al. 2001; Nagata et al. 2004; Hsu et al. 2010; Hoedjes et al. 2011; Srinivasan et al. 2012). The virus is transmitted by both larvae and adults, but only larvae can acquire the virus from infected plants. Virus transmission is in a persistent propagative manner: once a thrips has acquired the virus, it can transmit the virus for the remainder of its lifetime (Nagata et al. 1999; Kumar and Rawal 1999). The virus is mechanically sap-transmissible, and identification of the virus is confirmed using saptransmission to Nicotiana benthamiana (Cortes et al. 1998; Schwartz and Mohan 2008). The virus is not seed-borne and appears to be found in the bulbs. Likewise, the virus does not survive in soil (Mohan and Moyer 2004; du Toit et al. 2004, 2007). Virion properties and genome The virions are spherical and membrane bound 80–120 nm in diameter (Cortes et al. 1998). The genome comprises three unique molecules of negative or ambisense ssRNA, designated as L (large) 8880 nt (FJ623474 = NC_029799), M (medium) 4838 nt (AF214014 = NC_039232), and S (small) 3105 nt (AF001387 = NC_029800) (Tomassoli et al. 2009; Bag et al. 2009, 2010, 2012, 2015).

Leek yellow stripe virus Taxonomic position Genus: Potyvirus

(LYSV)

Family: Potyviridae

LYSV infection in plants of Allium cepa was reported from Iran and Turkey (Shahraeen et al. 1998, 2008; Fidan et al. 2013; Sevik and Akcura 2013). The virus-infected onion plants exhibit yellow stripe symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of LYSV, refer to Allium ampeloprasum var. porrum.

Onion yellow dwarf virus Taxonomic position Genus: Potyvirus

(OYDV)

Family: Potyviridae

Geographical distribution OYDV was first described by Melhus et al. (1929). The virus is probably distributed worldwide in onion crops in countries such as France, Morocco, Spain, Egypt, Greece, Iran, Italy, Japan, Turkey, China, Ecuador, India, and the former Yugoslavia (Conci et al. 1999; Hoa et al. 2003; Arya et al. 2006; Shahraeen et al. 2008; Elnagar et al. 2009; Fidan et al. 2009; Ward et al. 2009a; Kumar et al. 2011, 2012; Naderi Saffar et al. 2013; Sevik and Akcura 2013; Manglli et al. 2014; Sivaprasad et al. 2017). Symptoms and host(s) The first symptoms of OYDV in young onions are yellow streaks at the bases of the first true leaves. All leaves developing after these initial symptoms show symptoms ranging from yellow streaks to complete yellowing of leaves. Leaves are sometimes crinkled and flattened and tend to fall over. Scapes

Allium cepa (Onion)

61

may show extensive yellowing, twisting, and curling with small flower heads and poor quality seed. Bulbs are undersized. The natural host range of this virus is restricted to the genus Allium, mainly onion and garlic. Transmission The virus is transmitted by 60 aphid species, including Myzus persicae, in a non-persistent manner. The virus is transmitted by mechanical sap-inoculation, and is restricted to Allium spp. Leek does not appear to be affected. The virus is not transmissible by contact between plants, not transmissible by seed, and not transmissible by pollen. Virion properties and genome The virions are non-enveloped and flexuous filaments 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense, single-stranded RNA of 10,538 nt (AJ510223 = NC_005029) (Chen et al. 2003; Conci et al. 1999; Hoa et al. 2003; Sharma et al. 2014; Revers and Garcia 2015; Wylie et al. 2017).

Shallot latent virus Taxonomic position Genus: Carlavirus

(SLV)

Family: Betaflexiviridae

SLV infection in plants of Allium cepa was reported from the Netherlands and Turkey (Bos et al. 1978; Fidan et al. 2009; Sevik and Akcura 2013). The virus-infected onion plants exhibit symptomless infections. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SLV, refer to Allium cepa var. aggregatum.

Shallot virus X

(ShVX)

Taxonomic position Genus: Allexivirus

Family: Alphaflexiviridae

ShVX infection in plants of Allium cepa was reported from Sudan (Hamed et al. 2012a). The virusinfected onion plants exhibit mild mottling symptoms. The virus is transmitted by a mite vector, Aceria tulipae, and also by mechanical sap-inoculation. The use of infected vegetative propagative material is another source of virus spread. For more details of ShVX, refer to Allium cepa var. aggregatum.

Sint-Jan onion latent virus Taxonomic position Genus: Carlavirus

(SJOLV)

Family: Betaflexiviridae

Geographical distribution SJOLV infection was first reported in plants of Allium spp. from the Netherlands by Van Dijk (1993).

A

62

Allium cepa (Onion)

Symptoms and host(s) The virus-infected onion plants do not exhibit any symptoms. Transmission The virus is transmitted by an aphid vector, Myzus ascalonicus, in a non-persistent manner. The virus is also transmitted by mechanical sap-inoculation to less than three families. Virion properties and genome The virions are flexuous filaments about 650 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Adams et al. 2004).

Tobacco rattle virus

(TRV)

Taxonomic position Genus: Tobravirus

Family: Virgaviridae

TRV infection in plants of Allium cepa was reported from Egypt (Hamed et al. 2012b). The virusinfected onion plants exhibit symptoms of yellowing, malformation, yellow striping, white necrotic stripes, and stunting. The virus is transmitted by nematode vectors and also by mechanical sapinoculation to a large number of host plants. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Allium cepa was reported from India (Sivaprasad et al. 2010; Asadhi et al. 2016). The virus-infected onion plants exhibit symptoms of straw-colored, irregular, necrotic lesions on the young leaves and flower stalks. A drastic reduction in bulb size, necrosis of bulbs, wilting, and necrosis of plants leads to a reduction in yield. The virus is transmitted by the thrips vectors (Thrips tabaci and Frankliniella occidentalis), the virus present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible. The virus is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato black ring virus Taxonomic position Genus: Nepovirus

(TBRV)

Family: Secoviridae

Allium cepa (Onion)

63

TBRV infection in plants of Allium cepa was reported from Northern Ireland (Calvert and Harrison 1963). The virus-infected onion plants exhibit symptoms of irregular longitudinal chlorotic markings and stunting. The virus is transmitted by the nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TBRV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Allium cepa was reported from Italy, Brazil, Serbia, and Georgia (Mullis et al. 2004; Stankovic et al. 2012). The virus-infected onion plants exhibit symptoms of tip dieback, necrotic lesions, and chlorosis. The virus is transmitted by thrips vectors in a persistent propagative manner, and also through sap-inoculation, but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

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Arch Phytopathol Plant Protect 47:906–918 Sivaprasad Y, Bhaskara Reddy BV, Rekha Rani K, Raja Reddy K, Sai Gopal DVR (2010) First report of Tobacco streak ilarvirus infecting onion (Allium cepa). New Dis Rep 22:17 Sivaprasad Y, Garrido P, Mendez K, Pachacama S, Garrido A, Ramos L (2017) First report of Onion yellow dwarf virus infecting onion in Ecuador. J Plant Pathol 99:296 Srinivasan R, Sundaraj S, Pappu HR, Diffie S, Riley DG, Gitaitis RD (2012) Transmission of Iris yellow spot virus by Frankliniella fusca and Thrips tabaci (Thysanoptera: Thripidae). J Econ Entomol 105:40–47 Stankovic I, Bulajic A, Vucurovic A, Ristic D, Miojevic K, Nikolic D, Krstic B (2012) First report of Tomato spotted wilt virus infecting onion and garlic in Serbia. Plant Dis 96:918 Sujitha A, Bhaskara Reddy BV, Sivaprasad Y, Usha R, Sai Gopal DVR (2012) First report of Groundnut bud necrosis virus infecting onion (Allium cepa). Aust Plant Dis Notes 7:183–187 Tomassoli L, Tiberini A, Masenga V, Vicchi V, Turina M (2009) Characterization of Iris yellow spot virus isolates from onion crops in Northern Italy. J Plant Pathol 91:733–739 Trkulja V, Mihic J, Kovacic D, Stankovic I, Bulajic A, Vucurovic A, Krstic B (2013) First report of Iris yellow spot virus infecting onion in Bosnia and Herzegovina. Plant Dis 97:430 Van Dijk P (1993) Carlavirus isolates from cultivated Allium species represent three viruses. Neth J Plant Pathol 99:233–257 Velásquez-Valle R, Reveles-Torre LR, Salas-Muñez S, Mauricio-Castillo JA, Pappu HR (2016) First confirmed report of Iris yellow spot virus in onion nurseries in Zacatecas, Mexico. Plant Dis 100:1509 Ward LI, Perez-Egusquiza Z, Fletcher JD, Clover GRG (2009a) A survey of viral diseases of allium crops in New Zealand. Australas Plant Pathol 38:533–539 Ward LI, Perez-Egusquiza Z, Fletcher JD, Ochoa Corona FM, Tang JZ, Liefting LW, Martin EJ, Quinn BD, Pappu HR, Clover GRG (2009b) First report of Iris yellow spot virus on Allium cepa in New Zealand. Plant Pathol 58:406 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

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Allium cepa var. aggregatum (Shallot)

Allium cepa var. aggregatum (Shallot) Family: Amaryllidaceae

Garlic virus B Taxonomic position Genus: Allexivirus

Spice crop

(GarV-B)

Family: Alphaflexiviridae

GarV-B infection in plants of Allium cepa var. aggregatum was reported from New Zealand (Ward et al. 2009). The virus-infected shallot plants exhibit chlorotic streaking symptoms. The virus is transmitted by mite vectors and also by mechanical sap-inoculation. The use of infected vegetative propagative material is another source of virus spread. For more details of GarV-B, refer to Allium sativum.

Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

IYSV infection in plants of Allium cepa var. aggregatum was reported from France and Zimbabwe (Huchette et al. 2008; Karavina and Gubba 2017). The leaves of the virus-infected shallot plants exhibit symptoms of numerous necrotic, diamond-shaped lesions. The virus is transmitted by thrips vectors in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of IYSV, refer to Iris spp. or Allium cepa.

Onion yellow dwarf virus Taxonomic position Genus: Potyvirus

(OYDV)

Family: Potyviridae

OYDV infection in plants of Allium cepa var. aggregatum is cosmopolitan in distribution (Ward et al. 2009; Naderi Saffar et al. 2013; Milosevic et al. 2015). The virus-infected shallot plant leaves show irregular yellow striping to almost complete yellowing and also downward curling, flattening, and crinkling. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of OYDV, refer to Allium cepa.

Shallot latent virus Taxonomic position Genus: Carlavirus

(SLV)

Family: Betaflexiviridae

Allium cepa var. aggregatum (Shallot)

67

Geographical distribution SLV infection in plants of Allium cepa var. aggregatum is distributed worldwide (van Dijk 1993b; Barg et al. 1997; Ward et al. 2009). Symptoms and host(s) The virus-infected shallot plants do not exhibit any symptoms. The natural host range of this virus is restricted to members of the family Amaryllidaceae. This includes species of the genus Allium, such as onion, leek, garlic, shallot, Α. scorodoprasum, A. cepa var. perutile, A. cepa var. agregatum, A. ampeloprasum var. sectivum, A. chinense, and A. fistulosum. Transmission The virus is transmitted by aphid vectors such as Aulacorthum solani, Myzus ascolonicus, M. persicae, Neutoxoptera formosana, Aphis gossypii, and A. fabae in a non-persistent manner. The virus is also transmitted by mechanical sap-inoculation to a number of Amaryllidaceae members, and local lesions are produced on Chenopodium spp., Celosia argentea, and Vicia faba (van Dijk 1993b). The virus is mainly disseminated through the use of virus-infected vegetative propagative materials and not through true seed. Virion properties and genome The virions are flexuous filaments about 650–652 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8363 nt (AJ292226 = NC_003557) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Bos et al. 1978; Adams et al. 2004).

Shallot virus X

(ShVX)

Synonyms Shallot mite-borne latent virus (SMbLV) Taxonomic position Genus: Allexivirus

Family: Alphaflexiviridae

Geographical distribution ShVX infection in plants of Allium cepa var. aggregatum was initially identified and described in Russia and also observed in the India, the Netherlands, Germany, Italy, New Zealand, China, and Thailand, probably occurring worldwide (Van Dijk et al. 1991; Van Dijk and Van Der Vlugt 1994; Majumder et al. 2008; Perez-Egusquiza et al. 2009; Taglienti et al. 2015). Symptoms and host(s) The virus-infected shallot leaves exhibit mild mosaic and chlorotic symptoms. The host range is extremely narrow and ShVX has been experimentally transmitted to onion, where systemic infection was symptomless, and to Chenopodium murale, which reacted with local lesions. Transmission The virus is transmitted by a mite vector, Aceria tulipae, and also the virus is transmitted by mechanical sap-inoculation. The use of infected vegetative propagative material is another source of virus spread.

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68

Allium cepa var. aggregatum (Shallot)

Virion properties and genome The virions are very flexuous filaments about 800 nm in length and 12 nm in diameter. The genome consists of a single molecule of single-stranded RNA of 8832 nt (M97264 = NC_003795) and comprises six ORFs, encoding, in order, the replication-related proteins, the first two proteins of a TGB, a serine-rich protein of unknown function, the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 28 kDa in size (Kanyuka et al. 1992; Vishnichenko et al. 1993; Adams et al. 2004).

Shallot yellow stripe virus Taxonomic position Genus: Potyvirus

(SYSV)

Family: Potyviridae

Geographical distribution SYSV infection in plants of Allium cepa var. aggregatum was reported from Asia and the Netherlands (Van Dijk 1993a; Chittarath et al. 2017). Symptoms and host(s) The virus-infected shallot plants exhibit symptoms of mild striping on young leaves. Transmission The virus is transmitted by aphid vectors, Acyrthosiphon pisum and Myzus cymbalariae, in a nonpersistent manner. The virus is transmitted by mechanical sap-inoculation (Van Dijk 1993a). The use of infected planting material is the primary mode of virus spread. Virion properties and genome The virions are non-enveloped and flexuous filaments 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 10429 nt (AJ865076 = NC_007433) (Chen et al. 2005; Revers and Garcia 2015; Wylie et al. 2017).

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Bos L, Huttinga H, Maat DZ (1978) Shallot latent virus a new Carlavirus. Neth J Plant Pathol 84:227–237 Chen J, Wei CB, Zheng HY, Shi YH, Adams MJ, Lin L, Zhang QY, Wang SJ, Chen JP (2005) Characterisation of the welsh onion isolate of Shallot yellow stripe virus from China. Arch Virol 150(10):2091–2099 Chittarath K, Rungsawang W, Pongsapich P, Kong GA, Thomas JE, Geering ADW (2017) First records of the potyviruses Chilli ringspot virus and Shallot yellow stripe virus from Laos. Aust Plant Dis Notes 12:53 de Barg EL, Vetten HJ, Green SK (1997) Viruses of alliums and their distribution in different Allium crops and geographical regions. Acta Hortic 433:607–616 Huchette O, Bellamy C, Filomenko R, Pouleau B, Seddas S, Pappu HR (2008) Iris yellow spot virus on shallot and onion in France. Plant Health Prog. Online. https://doi.org/10.10194/PHP-2008-0610-01-BR Kanyuka KV, Vishnichenko VK, Levay KE, Kondrikov DY, Ryabov EV, Zavriev SK (1992) Nucleotide sequence of Shallot virus X RNA reveals a 50 - proximal cistron closely related to those of the potexviruses and a unique arrangement of the 30 -proximal cistrons. J Gen Virol 73:2553–2560 Karavina C, Gubba A (2017) Iris yellow spot virus in Zimbabwe: Incidence, severity and characterization of Alliuminfecting isolates. Crop Prot 94:69–76

Allium chinense (Chinese scallion, Rakkyo)

69

Majumder S, Arya M, Pant RP, Baranwal VK (2008) Shallot virus X in Indian shallot a new virus report for India. Plant Pathol 57:396 Milosevic D, Gvozdanovic-Varga J, Ignjatov M, Nikolic Z, Vucurovic I, Vucurovic A, Stankovic I (2015) First report of Onion yellow dwarf virus infecting shallot in Serbia. Plant Dis 99:1450 Naderi Saffar Z, Torabi S, Naghavi M, Golnaraghi AR, Aryakia E (2013) Onion yellow dwarf virus on leek, onion, shallot and welsh onion in Iran. J Plant Pathol 95:S4.73 Perez-Egusquiza Z, Ward LI, Clover GRG, Fletcher JD, Van Der Vlugt RAA (2009) First report of Shallot virus X in shallot in New Zealand. Plant Pathol 58:407 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Taglienti A, Taviani P, Paoletti S, Tomassoli L (2015) First report of Shallot virus X infecting shallot in Italy. New Dis Rep 32:28 Van Dijk P (1993a) Survey and characterization of potyviruses and their strains of Allium species. Neth J Plant Pathol 99:1–48 Van Dijk P (1993b) Carlavirus isolates from cultivated Allium species represent three viruses. Neth J Plant Pathol 99:233–257 Van Dijk P, Verbeek M, Bos L (1991) Mite-borne virus isolates from cultivated Allium species, and their classification into two new rymoviruses in the family Potyviridae. Neth J Plant Pathol 97:381–399 Van Dijk P, Van Der Vlugt RAA (1994) New mite-borne virus isolates from rakkyo, shallot and leek species. Eur J Plant Pathol 100:269–277 Vishnichenko VK, Konareva TN, Zavriev SK (1993) A new filamentous virus in shallot. Plant Pathol 42:121–126 Ward LI, Perez-Egusquiza Z, Fletcher JD, Clover GRG (2009) A survey of viral diseases of allium crops in New Zealand. Australas Plant Pathol 38:533–539 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Allium chinense (Chinese scallion, Rakkyo) Family: Amaryllidaceae

Spice crop

Asparagus virus 3

(AV-3)

Synonyms Scallion virus X (ScaVX) Taxonomic position Genus: Potexvirus

Family: Alphaflexiviridae

ScaVX infection in plants of Allium chinense was reported from China (Chen et al. 2002). The virus is transmitted by means not involving a vector. The virus is transmitted by mechanical sap-inoculation. Use of infected planting material is the primary source of virus spread. For more details of AV-3, refer to Asparagus officinalis.

Onion yellow dwarf virus Taxonomic position Genus: Potyvirus

(OYDV)

Family: Potyviridae

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Allium chinense (Chinese scallion, Rakkyo)

OYDV infection in plants of Allium chinense was reported from China (Sako et al. 1991a; Wu et al. 2009). The virus-infected rakkyo plants exhibit symptoms of mosaic, chlorotic streaking, twisting, and crinkling on the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of OYDV, refer to Allium cepa.

Scallion mosaic virus Taxonomic position Genus: Potyvirus

(ScaMV)

Family: Potyviridae

Geographical distribution ScaMV infection in plants of Allium chinense was reported from China (Chen et al. 2002). Symptoms and host(s) The virus-infected rakkyo plants exhibit mild stripe symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus could be transmitted mechanically to scallion but not to other Allium species (including garlic). Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense, single-stranded RNA 9324 nt (AJ316084 = NC_003399), encoding a 341.3 kDa polyprotein of 3001 amino acids (Chen et al. 2002; Revers and Garcia 2015; Wylie et al. 2017).

Shallot virus X

(ShVX)

Synonyms Shallot mite-borne latent virus (SMbLV) Taxonomic position Genus: Allexivirus

Family: Alphaflexiviridae

ShVX infection in plants of Allium chinense was reported from the Netherlands (Van Dijk and Van Der Vlugt 1994). The virus is transmitted by a mite vector, Aceria tulipae, and also through mechanical sapinoculation. The use of infected vegetative propagative material is another source of virus spread. For more details of ShVX, refer to Allium cepa var. aggregatum.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

Allium fistulosum (Green onion, Welsh onion)

71

TMV infection in plants of Allium chinense was reported from Japan (Nagata and Sako 1989; Sako et al. 1991b; Kwon and Sako 1994). The virus-infected rakkyo plants exhibit symptoms of stunting and slight twisting of the leaves. There is no known vector for this virus. The virus is mechanically sap-transmissible and also through contact between plants. For more details of TMV, refer to Nicotiana tabacum.

References Chen J, Zheng HY, Chen JP, Adams MJ (2002) Characterisation of a Potyvirus and a Potexvirus from Chinese scallion. Arch Virol 147(4):683–693 Kwon S-B, Sako N (1994) A new strain of Tobacco mosaic virus infecting rakkyo (Allium chinense G. Don). Ann Phytopath Soc Jpn 60:36–44 Nagata R, Sako N (1989) Garlic latent virus and Tobacco mosaic virus detected in rakkyo. Ann Phytopath Soc Japan 55:537 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Sako I, Nakasone W, Okada K, Ohki ST, Osaki T, Inouye T (1991a) Yellow streak of rakkyo (Allium chinense G. Don), a newly recognized disease caused by Garlic latent virus and Onion yellow dwarf virus. Ann Phytopathol Soc Jpn 57:65–69 Sako I, Osaki T, Nakasone W, Inouye T (1991b) Yellow mosaic of rakkyo (Allium chinense G. Don.) caused by Tobacco mosaic virus and Garlic latent virus, and its occurrence in the field. Proc Kansai Plant Protect Soc 33:21–28 Van Dijk P, Van Der Vlugt RAA (1994) New mite-borne virus isolates from rakkyo, shallot and leek species. Eur J Plant Pathol 100:269–277 Wu CC, Hong N, Liu Y, Xu WX, Wang GP (2009) First report of Onion yellow dwarf virus in Allium chinense in China. Plant Dis 93:761 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Allium fistulosum (Green onion, Welsh onion) Family: Amaryllidaceae

Garlic virus D Taxonomic position Genus: Allexivirus

Spice crop

(GarV-D)

Family: Alphaflexiviridae

GarV-D infection in plants of Allium fistulosum was reported from Poland (Paduch-Cichal and Bereda 2017). The virus is transmitted by a mite vector, Aceria tulipae, and also by mechanical sap-inoculation. The use of infected vegetative propagative material is another source of virus spread. For more details of GarV-D, refer to Allium sativum.

Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

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Allium fistulosum (Green onion, Welsh onion)

IYSV infection in plants of Allium fistulosum was reported from Indonesia and the United States (Pappu and Rauf 2013; Tabassum et al. 2016). The virus-infected welsh onion plants exhibit symptoms consisting of elliptical to spindle-shaped, straw-colored, irregular, chlorotic lesions with occasional green islands on the leaves. The virus is transmitted by a thrips vector, Thrips tabaci, in a persistent propagative manner, and the virus is also mechanically sap-transmissible. For more details of IYSV, refer to Iris spp. or Allium cepa.

Onion yellow dwarf virus Taxonomic position Genus: Potyvirus

(OYDV)

Family: Potyviridae

OYDV infection in plants of Allium fistulosum was reported from Iran and Japan (Fukami et al. 1989; Fukami and Ishii 1991; Sako et al. 1994; Naderi Saffar et al. 2013). The virus-infected welsh onion plants exhibit symptoms of mosaic and yellow streak striping, curling, and distortion of flower stems. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of OYDV, refer to Allium cepa.

Shallot yellow stripe virus (Welsh onion yellow stripe virus) Taxonomic position Genus: Potyvirus

(SYSV)

Family: Potyviridae

SYSV infection in plants of Allium fistulosum was reported from Brazil, Indonesia, China, and Japan (Liu et al. 1996; van der Vlugt et al. 1999; Chen et al. 2005). The virus-infected welsh onion plants exhibit yellow leaf striping symptoms. The virus is transmitted by aphid vectors, Myzus persicae, Rhopalosiphum maidis, and Acyrthosiphon pisum, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SYSV, refer to Allium cepa var. aggregatum.

References Chen J, Wei C-B, Zheng H-Y, Shi Y-H, Adams MJ, Lin L, Zhang Q-Y, Wang S-J, Chen J-P (2005) Characterisation of the welsh onion isolate of Shallot yellow stripe virus from China. Arch Virol 150:2091–2099 Fukami M, Ishii K (1991) Occurrence of Garlic latent virus (GLV) and Onion yellow dwarf virus (OYDV) on seedless Welsh onions “Bozushirazu” in Chiba Prefecture, and influence of their infection on their yield. Bull Chiba Pferectural Agril Experiment Station 32:9–17 Fukami M, Natsuaki KT, Tomaru K (1989) Detection of Onion yellow dwarf virus in Allium fistulosum by gelatin particle agglutination test. Ann Phytopathol Soc Jpn 55:542 Liu HM, Zhang QP, Li JR (1996) Studies on the characteristics of Welsh-onion yellow stripe virus. J Shandong Agric Univ 27:303–310 Naderi Saffar Z, Torabi S, Naghavi M, Golnaraghi AR, Aryakia E (2013) Onion yellow dwarf virus on leek, onion and welsh onion in Iran. J Plant Pathol 95:S4.73 Paduch-Cichal M, Bereda M (2017) Viruses infecting ornamental allium species in Poland. J Plant Pathol 99(2). https:// doi.org/10.4454/jpp.v99i2.3882 Pappu HR, Rauf A (2013) First report of Iris yellow spot virus infecting green onion in Indonesia. Plant Dis 97:1665 Sako I, Takami T, Nakasone W, Osaki T, Inouye T (1994) Occurrence of Garlic latent virus and Onion yellow dwarf virus in seedless Welsh onion (cv. Bozushirazu) and influence of virus reinfection on their yield. Proc Kansai Plant Protect Soc 36:21–27

Allium sativum (Garlic)

73

Tabassum A, Reitz S, Rogers P, Pappu HR (2016) First report of Iris yellow spot virus infecting green onion (Allium fistulosum) in the United States. Plant Dis 100:2539 van der Vlugt RAA, Steffens P, Cuperus C, Barg E, Lesemann D-E, Bos L, Vetten HJ (1999) Further evidence that Shallot yellow stripe virus (SYSV) is a distinct potyvirus and reidentification of Welsh onion yellow stripe virus as a SYSV strain. Phytopathology 89:148–155

Allium macrostemon (Japanese garlic) Family: Amaryllidaceae

Spice crop

Scallion mosaic virus Taxonomic position Genus: Potyvirus

(ScaMV)

Family: Potyviridae

ScaMV infection in plants of Allium macrostemon was reported from Japan (Oshima et al. 2016). The virus-infected Japanese garlic plants exhibit symptoms of the mild stripping of leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ScaMV, refer to Allium chinense.

References Oshima K, Muraoka S, Yasaka R, Adachi S, Tokuda M (2016) First report of Scallion mosaic virus on wild Japanese garlic (Allium macrostemon) in Japan. J Gen Plant Pathol 82:61–64

Allium sativum (Garlic) Family: Amaryllidaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Spice crop

(CMV)

Family: Bromoviridae

CMV infection in plants of Allium sativum was reported from Yugoslavia (Stefanac 1980). The virusinfected garlic plants exhibit stunting symptoms. The virus is non-persistently transmitted by more than 86 spp. of aphid vectors Aphis gossypii, A. craccivora, Acyrthosiphon pisum, and Myzus persicae which are the major vectors. The virus is transmitted by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

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Allium sativum (Garlic)

Garlic common latent virus Taxonomic position Genus: Carlavirus

(GarCLV)

Family: Betaflexiviridae

Geographical distribution GarCLV is detected in most of the Allium sativum-growing countries, including Southeast Asia, Australia, Turkey, Israel, Sudan, Brazil, North Africa (Sudan), Europe, the USA, Nigeria, and China (Delecolle and Lot 1981; Barg et al. 1994; Pappu et al. 2005; Gieck et al. 2007; Klukackova et al. 2007; Mohammad et al. 2007; Fidan and Baloglu 2009b; Majumder and Baranwal 2009, 2014; Fayad-Andre et al. 2011; Parrano et al. 2012; Pramesh et al. 2012; Wylie et al. 2012; Hamed et al. 2013; Majumder and Johari 2014; Vucurovic et al. 2015; Majumder et al. 2016; Voncina et al. 2016). Symptoms and host(s) The virus causes either mild to severe mosaic symptoms or no symptoms in singly infected garlic plants depending on the cultivar and environmental factors. However, in Sudan certain virus-infected garlic cultivars exhibited mosaic and/or mottling symptoms (Bellardi et al. 1995; Hamed et al. 2013). Transmission The virus is transmitted by aphid vectors in a non-persistent manner (Van Dijk 1993; Barg et al. 1994). The virus is also mechanically sap-transmissible with difficulty to Allium, Solanaceae, and Chenopodiaceae members. The virus is not transmitted through true seed, but vegetative propagation is the prime mode of virus spread. Virion properties and genome The virions are flexuous filaments particles and measure 650 nm length and 12–15 nm in diameter. The genome is a monopartite, linear, positive-sense, single-stranded RNA of 8638 nt (JF320810 = NC_016440) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Tsuneyoshi et al. 1998; Fajardo et al. 2001; Adams et al. 2004; Wylie et al. 2012; Pramesh and Baranwal 2013).

Garlic dwarf virus

(GDV)

Taxonomic position Genus: Fijivirus

Family: Reoviridae

Geographical distribution GDV infection in plants of Allium sativum was reported in the Eurasian region (France) (Lot et al. 1994). Symptoms and host(s) The leaves of virus-infected garlic plants assume a darker green color with swollen veins and enations. The initial symptoms are red tips of the basal leaves. The majority of the affected plants do not develop normally, showing a “tulip” or “fan” appearance with all leaves originating from the same point, and sometimes severe dwarfing is noticed. The bulbs from the diseased plants often appear pear-shaped, spongy, and wrinkled.

Allium sativum (Garlic)

75

Transmission There is no known vector for this virus. The primary spread of the virus takes place by the use of infected vegetative propagative planting material. The virus is not mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, icosahedral with a double capsid structure, about 65–70 nm in diameter. Pentameric turrets sit on the outside of the innermost capsid. The outer capsid has a T = 13 icosahedral symmetry and the inner capsid a T = 2 icosahedral symmetry. The genome is linear double-stranded RNA containing 10 segments coding for 12 proteins (Francki and Boccardo 1983; Lot et al. 1994; Harding and Dale 2011).

Garlic mite-borne filamentous virus Taxonomic position Genus: Allexivirus

(GarMbFV)

Family: Alphaflexiviridae

Geographical distribution GarMbFV infection in plants of Allium sativum was reported from Brazil, Poland, and Japan (Yamashita et al. 1996; Melo-Filho et al. 2004; Torres et al. 2004; Fayad-Andre et al. 2011; Bereda and Paduch-Cichal 2016; Bereda et al. 2017). Symptoms and host(s) The virus-infected garlic plants exhibit systemic symptoms. Transmission The virus is transmitted by a mite vector, Aceria tulipae. The virus is mechanically sap-transmissible. The use of infected vegetative propagative material is another source of virus spread. Virion properties and genome The virions are very flexuous filaments about 800 nm in length and 12 nm in diameter. The genome consists of a single molecule of linear single-stranded RNA. A partial coat protein sequence of 762 nt is available (X98991). The genome is comprised of six ORFs, encoding, in order, the replication-related proteins, the first two proteins of a TGB, a serine-rich protein of unknown function, the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 28 kDa in size (Adams et al. 2004).

Garlic virus A Taxonomic position Genus: Allexivirus

(GarV-A)

Family: Alphaflexiviridae

Geographical distribution GarV-A infection in plants of Allium sativum was reported from Australia, India, Brazil, Poland, Korea, and Argentina (Lunello et al. 2000; Koo et al. 2002; Melo-Filho et al. 2004; Perotto et al. 2010; Chodorska et al. 2012; Gawande et al. 2015; Bereda and Paduch-Cichal 2016; Bereda et al. 2017).

A

76

Allium sativum (Garlic)

Symptoms and host(s) The virus-infected garlic plants exhibit mild mosaic-like symptoms on the leaves. Transmission The virus is transmitted by a mite vector, Aceria tulipae (Kang et al. 2007). The virus is mechanically sap-transmissible. The use of infected vegetative propagative material is another source of virus spread. Virion properties and genome The virions are flexible filamentous, about 800 nm in length and 12 nm in diameter. The genome consists of linear single-stranded RNA of 8660 nt (AB010300 = NC_003375) and comprises six ORFs, encoding, in order, the replication-related proteins, the first two proteins of a TGB, a serine-rich protein of unknown function, the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 28 kDa in size (Sumi et al. 1993, 1999; Adams et al. 2004; Wylie et al. 2012).

Garlic virus B Taxonomic position Genus: Allexivirus

(GarV-B)

Family: Alphaflexiviridae

Geographical distribution GarV-B infection in plants of Allium sativum was reported from Poland, Finland, Brazil, Iran, Italy, and the US Pacific Northwest (Sumi et al. 1993; Melo-Filho et al. 2004; Shahraeen et al. 2008; Gieck et al. 2009; Chodorska et al. 2012; Bereda and Paduch-Cichal 2016; Bereda et al. 2017; Taglienti et al. 2018). Symptoms and host(s) The virus-infected garlic plants exhibit yellowing and stunting symptoms. Transmission The virus is transmitted by a mite vector, Aceria tulipae (Kang et al. 2007). The virus is mechanically sap-transmissible. The use of infected vegetative propagative material is another source of virus spread. Virion properties and genome The virions are flexible filamentous, about 800 nm in length and 12 nm in diameter. The genome consists of linear single-stranded RNA of 8336 nt (KM379144 = NC_025789) and comprises six ORFs, encoding, in order, the replication-related proteins, the first two proteins of a TGB, a serine-rich protein of unknown function, the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 28 kDa in size (Sumi et al. 1993; Adams et al. 2004; Wylie et al. 2012).

Garlic virus C Taxonomic position Genus: Allexivirus

(GarV-C)

Family: Alphaflexiviridae

Allium sativum (Garlic)

77

Geographical distribution GarV-C infection in plants of Allium sativum was reported from Poland, Finland, New Zealand, Australia, and Brazil (Sumi et al. 1999; Melo-Filho et al. 2004; Torres et al. 2004; Shahraeen et al. 2008; Ward et al. 2009; Perotto et al. 2010; Fayad-Andre et al. 2011; Chodorska et al. 2012; Wylie et al. 2012; Bereda and Paduch-Cichal 2016; Bereda et al. 2017). Symptoms and host(s) The virus-infected garlic plants exhibit yellowing and stunting symptoms. Transmission The virus is transmitted by a mite vector, Aceria tulipae (Kang et al. 2007). The virus is mechanically sap-transmissible. The use of infected vegetative propagative material is another source of virus spread. Virion properties and genome The virions are flexible filamentous, about 800 nm in length and 12 nm in diameter. The genome consists of linear single-stranded RNA of 8405 nt (AB010302 = NC_003376) and comprises six ORFs, encoding, in order, the replication-related proteins, the first two proteins of a TGB, a serine-rich protein of unknown function, the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 28 kDa in size (Sumi et al. 1999; Adams et al. 2004; Wylie et al. 2012).

Garlic virus D Taxonomic position Genus: Allexivirus

(GarV-D)

Family: Alphaflexiviridae

Geographical distribution GarV-D infection in plants of Allium sativum was reported from Brazil, New Zealand, Iran, Italy, Poland, Australia, India, and the US Pacific Northwest (Koo et al. 2002; Chen et al. 2004; Melo-Filho et al. 2004; Torres et al. 2004; Shahraeen et al. 2008; Gieck et al. 2009; Ward et al. 2009; Fayad-Andre et al. 2011; Wylie et al. 2012; Chodorska et al. 2013; Bereda et al. 2015, 2017; Bereda and PaduchCichal 2016; Khan et al. 2016; Taglienti et al. 2018). Symptoms and host(s) The virus-infected garlic plants exhibit symptoms of mosaic and chlorotic leaves Transmission The virus is transmitted by a mite vector, Aceria tulipae (Kang et al. 2007). The virus is mechanically sap-transmissible. The use of infected vegetative propagative material is another source of virus spread. Virion properties and genome The virions are flexible filamentous, about 800 nm in length and 12 nm in diameter. The genome consists of linear positive-sense single-stranded RNA of 8424 nt (KF555653 = NC_022961) and comprises six ORFs, encoding, in order, the replication-related proteins, the first two proteins of a TGB, a serine-rich protein of unknown function, the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 28 kDa in size (Sumi et al. 1993; Adams et al. 2004; Wylie et al. 2012).

A

78

Garlic virus E Taxonomic position Genus: Allexivirus

Allium sativum (Garlic)

(GarV-E)

Family: Alphaflexiviridae

Geographical distribution GarV-E infection in plants of Allium sativum was reported from China and Poland (Chen et al. 2001; Chodorska et al. 2013; Bereda and Paduch-Cichal 2016). Symptoms and host(s) The virus-infected garlic plants exhibit mosaic, deformation, and yellow stripe symptoms. Transmission The virus is transmitted by a mite vector, Aceria tulipae (Kang et al. 2007). The virus is mechanically sap-transmissible. The use of infected vegetative propagative material is another source of virus spread. Virion properties and genome The virions are flexible filamentous, about 800 nm in length and 12 nm in diameter. The genome consists of a single molecule of linear single-stranded RNA of 8451 nt (AJ292230 = NC_004012) and comprises six ORFs, encoding, in order, the replication-related proteins, the first two proteins of a TGB, a serine-rich protein of unknown function, the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 28 kDa in size (Chen et al. 2001; Adams et al. 2004).

Garlic virus X Taxonomic position Genus: Allexivirus

(GarV-X)

Family: Alphaflexiviridae

Geographical distribution GarV-X infection in plants of Allium sativum was reported from Italy, Argentina, Brazil, the USA, China, Japan, Korea, Poland, and India (Song et al. 1997; Chen et al. 2004; Baranwal et al. 2011; Chodorska et al. 2013; Parrano et al. 2012; Oliveira et al. 2014; Bereda and Paduch-Cichal 2016; Bereda et al. 2017; Taglienti et al. 2018). Symptoms and host(s) The virus-infected garlic plants exhibit mosaic leaf symptoms and stunting of the plants. Transmission The virus is transmitted by the mite vector, Aceria tulipae (Kang et al. 2007). The virus is mechanically sap-transmissible and is also transmitted through the cloves collected from virus-infected plants. Virion properties and genome The virions are flexuous and rod-shaped with a length of about 750 nm and width of 12 nm. The genome is single-stranded positive-sense RNA of 8106 nt (U89243 = NC_001800) and comprises six ORFs, encoding, in order, the replication-related proteins, the first two proteins of a TGB, a serine-rich protein of unknown function, the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 28 kDa in size (Song et al. 1997, 1998; Adams et al. 2004).

Allium sativum (Garlic)

79

Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

A

Family: Tospoviridae

IYSV infection in plants of Allium sativum was reported from the USA, Europe, Zimbabwe, and Asia (Bag et al. 2009; Gawande et al. 2010; Karavina et al. 2016; Karavina and Gubba 2017). The virusinfected garlic plants show straw-colored, diamond-shaped/spindle-shaped spots with poorly defined ends of the leaves. The spots coalesce to form larger patches on the leaves and are clearly visible on older leaves. In some cases dry, chlorotic and necrotic, oval- and diamond-shaped lesions are noticed on leaves and flower stalks. The virus is transmitted by the onion thrips Thrips tabaci in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of IYSV, refer to Iris spp. or Allium cepa.

Leek yellow stripe virus

(LYSV)

Synonyms Garlic mosaic virus, Garlic yellow streak virus

Taxonomic position Genus: Potyvirus

Family: Potyviridae

LYSV infection in plants of Allium sativum was reported from South and Southeast Asia, France, Israel, Venezuela, France, Java, Australia, Italy, China, Turkey, Syria, North America, Argentina, Brazil, Mexico, Japan, the Czech Republic, Ecuador, Croatia, Poland, and New Zealand (Mohamed and Young 1981; Dewan et al. 1995; Yamashita et al. 1995; Pappu et al. 2005; Navratil et al. 2006; Perez-Moreno et al. 2006; Wei et al. 2006; Gieck et al. 2007; Klukackova et al. 2007; Parrano et al. 2012; Fidan et al. 2013; Gupta et al. 2013; Chodorska et al. 2014b; Gawande et al. 2014; Testen et al. 2014; Oleas and Arahana 2016; Voncina et al. 2016). The virus-infected garlic plants exhibit yellow streak symptoms and the plants are stunted. The virus is transmitted by a number of aphid spp. in a non-persistent manner, and also by mechanical sap-inoculation. For more details of LYSV, refer to Allium ampeloprasum var. porrum.

Lettuce necrotic yellows cytorhabdovirus Taxonomic position Genus: Cytorhabdovirus

(LNYV)

Family: Rhabdoviridae

LNYV infection in plants of Allium sativum was reported from Victoria (Australia) (Sward 1990). The virus-infected garlic plants do not exhibit any specific symptoms. The virus is transmitted by an aphid vector, Hyperomyzus lactucae, in a persistent, propagative manner, and also by mechanical sapinoculation. For more details of LNYV, refer to Lactuca sativa.

80

Allium sativum (Garlic)

Onion yellow dwarf virus Taxonomic position Genus: Potyvirus

(OYDV)

Family: Potyviridae

OYDV infection in plants of Allium sativum is reported worldwide, including France, Italy, Syria, Turkey, China, Argentina, Japan, Canada, India, Egypt, Nepal, the Czech Republic, Ecuador, Croatia, and the USA (Lot et al. 1998; Stobbs and Van Driel 1999; Shiboleth et al. 2001; Pappu et al. 2005; Arya et al. 2006; Gieck et al. 2007; Klukackova et al. 2007; Mahmoud et al. 2007; Majumder et al. 2008, 2017; Elnagar et al. 2009; Fidan and Baloglu 2009a; Gawande et al. 2013; Majumder and Baranwal 2014; Majumder and Johari 2014; Oleas and Arahana 2016; Voncina et al. 2016). The virus on the garlic causes mild mosaic or very mild chlorotic stripes to bright yellow stripes, curling and distortion of flower stems, reduction in the number of flowers and seeds, and impairment of seed quality. The virus is transmitted by an aphid vector, Myzus persicae, in a non-persistent manner, and also by mechanical sapinoculation. The virus is not transmissible by contact between plants. For more details of OYDV, refer to Allium cepa.

Shallot latent virus Taxonomic position Genus: Carlavirus

(SLV)

Family: Betaflexiviridae

SLV infection in plants of Allium sativum was reported from Iran, China, Canada, Egypt, Greece, Syria, Slovenia, Italy, India, Argentina, the Netherlands, the Czech Republic, Ecuador, Poland, and Brazil (Van Dijk 1993; Dovas and Vovlas 2003; Klukackova et al. 2004; Mavric and Ravnikar 2005; Klukackova et al. 2007; Mohammad et al. 2007; Majumder et al. 2008; Arya et al. 2009; Torrico et al. 2010; Mituti et al. 2011; Chodorska et al. 2014a; Majumder and Baranwal 2014; Oleas and Arahana 2016). The virus-infected garlic plants do not exhibit any symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SLV, refer to Allium cepa var. aggregatum.

Shallot virus X

(ShVX)

Taxonomic position Genus: Allexivirus

Family: Alphaflexiviridae

ShVX infection in plants of Allium sativum was reported from Poland and Ecuador (Bereda and Paduch-Cichal 2016; Granda et al. 2017). The virus is transmitted by a mite vector, Aceria tulipae, and also by mechanical sap-inoculation. The use of infected vegetative propagative material is another source of virus spread. For more details of ShVX, refer to Allium cepa var. aggregatum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

Allium sativum (Garlic)

81

TSWV infection in plants of Allium sativum was reported from Serbia (Stankovic et al. 2012). The virus-infected garlic plants exhibit symptoms of chlorotic spots and streaks. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is not transmissible by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Arya M, Baranwal VK, Ahlawat YS, Singh L (2006) RT-PCR detection and molecular characterization of Onion yellow dwarf virus associated with garlic and onion. Curr Sci 91:1230–1234 Arya M, Majumder S, Baranwal VK (2009) Partial characterisation of coat protein gene of Shallot latent virus associated with garlic in India. Indian J Virol 20:9–11 Bag S, Rogers P, Watson R, Pappu HR (2009) First report of natural infection of garlic with Iris yellow spot virus in the United States. Plant Dis 93:839 Baranwal VK, Singh P, Jain RK, Joshi S (2011) First report of Garlic virus X infecting garlic in India. Plant Dis 95(9):1197 Barg E, Lesemann DE, Vetten HJ, Green SK (1994) Identification, partial characterization and distribution of viruses infecting Allium crops in South and Southeast Asia. Acta Hortic 358:251–258 Bellardi MG, Marani F, Betti L, Rabiti AL (1995) Detection of Garlic common latent virus (GCLV) in Allium sativum L. in Italy. Phytopathology 34:58–61 Bereda M, Paduch-Cichal E (2016) Allexiviruses – pathogens of garlic plants. Prog Plant Protect 56:302–311 Bereda M, Kalinowska E, Paduch-Cichal E, Stefan Szyndel M (2015) Low genetic diversity of a natural population of Garlic virus D from Poland. Eur J Plant Pathol 142:411–417 Bereda M, Paduch-Cichal E, Dabrowska E (2017) Occurrence and phylogenetic analysis of allexiviruses identified on garlic from China, Spain and Poland commercially available on the polish retail market. Eur J Plant Pathol 149:227–237 Chen J, Chen J, Adams MJ (2001) Molecular characterization of a complex mixture of viruses in garlic with mosaic symptoms in China. Arch Virol 146:1841–1853 Chen J, Zheng HY, Antoniw JF, Adams MJ, Chen JP, Lin L (2004) Detection and classification of allexiviruses from garlic in China. Arch Virol 149:435–445 Chodorska M, Nowak P, Szyndel MS, Paduch-Cichal E, Sala-Rejczak K (2012) First report of Garlic virus A, B and C in garlic in Poland. J Plant Pathol 94:S4.100 Chodorska M, Paduch-Cichal E, Szyndel MS, Kalinowska E (2013) First report of Garlic virus D, E and X on garlic in Poland. J Plant Pathol 95:S4.70 Chodorska M, Paduch-Cichal E, Kalinowska E, Szyndel MS (2014a) First report of Onion yellow dwarf virus, Garlic common latent virus and Shallot latent virus on Garlic in Poland. Plant Dis 98(6):858 Chodorska M, Paduch-Cichal E, Kalinowska E, Gaczkowska O, Lis M, Sierant B, Szyndel MS (2014b) First report of Leek yellow stripe virus in Foreign and Polish garlic plants in central Poland. J Plant Pathol 96:S4.120 Delecolle B, Lot H (1981) Viroses de l’ail: I. – Mise en évidence et essais de caracté risation par immuno électromicroscopie d’un complexe de trois virus chez différentes populations d’ail atteintes de mosaique. Agronomie 1:763–770 Dewan C, Pearson MN, Scheffer JJC (1995) Natural field infection of garlic by Garlic yellow streak virus in the Pukekohe area of New Zealand and associated problems with the introduction of new garlic cultivars. N Z J Crop Hortic Sci 23:97–102 Dovas CI, Vovlas C (2003) Viruses infecting Allium spp. in Southern Italy. J Plant Pathol 85:135 Elnagar S, El-Sheikh MAK, Abdel Wahab AS (2009) Effect of natural infection with Onion yellow dwarf virus (OYDV) on yield of onion and garlic crops in Egypt. In: 4th conference on recent technologies in agriculture, pp 34–39 Fajardo TVM, Nishijima M, Buso JA, Torres AC, Avila AC, Resende RO (2001) Garlic viral complex: identification of Potyvirus and Carlavirus in Central Brazil. Fitopatol Bras 26(3):619–626 Fayad-Andre MS, Dusi AN, Resende RO (2011) Spread of viruses in garlic fields cultivated under different agricultural production systems in Brazil. Trop Plant Pathol 36:341–349 Fidan H, Baloglu S (2009a) First report of Onion yellow dwarf virus and Leek yellow stripe virus in Garlic in Turkey. Plant Dis 93:672 Fidan H, Baloglu S (2009b) First report of Garlic common latent virus in garlic in turkey. J Plant Pathol 91(4 Suppl):S4.99

A

82

Allium sativum (Garlic)

Fidan H, Caglar BK, Baloglu S, Yilmaz MA (2013) Urginea maritima (L.) is a new host of Allexivirus group on onion and garlic plants in Turkey. In: Baktir I et al (eds) Proceedings of the XIth IS on flower bulbs and herbaceous perennials. Acta Hortic 1002:309–312 Francki RIB, Boccardo G (1983) The plant Reoviridae. In: Joklik WK (ed) The Reoviridae. Plenum Press, New York, pp 505–563 Gawande SJ, Khar A, Lawande KE (2010) First report of Iris yellow spot virus on garlic in India. Plant Dis 94:1066 Gawande SJ, Chimote KP, Gurav VS, Gopal J (2013) Distribution and natural incidence of Onion yellow dwarf virus (OYDV) on garlic and its related Allium species in India. Indian J Hortic 70:544–548 Gawande SJ, Gurav VS, Ingle AA, Gopal J (2014) First report of Leek yellow stripe virus in Allium sativum in Western India. Plant Dis 98:1015 Gawande SJ, Gurav VS, Ingle AA, Gopal J (2015) First report of Garlic virus A in Garlic From India. Plant Dis 99:1288 Gieck SL, Pappu HR, Hamm PB, David NL (2007) First report of Onion yellow dwarf virus, Leek yellow stripe virus, and Garlic common latent virus in Garlic in Oregon. Plant Dis 91:461 Gieck SL, Hamm PB, David NL, Pappu HR (2009) First report of Garlic virus B and Garlic virus D in Garlic in the Pacific Northwest. Plant Dis 93:431 Granda R, Landazuri G, Arkhipov AV (2017) First report of Shallot virus X in garlic in Ecuador. Plant Dis 101:1066 Gupta N, Prabha K, Islam S, Baranwal VK (2013) First report of Leek yellow stripe virus in garlic from India. J Plant Pathol 95:S4.69–S4.77 Hamed K, Menzel W, Mohamed ME, Bakheet KA, Winter S (2013) First report of Garlic common latent virus infecting garlic in Sudan. Plant Dis 97:562 Harding RM, Dale JL (2011) Fijivirus. Reoviridae. In: The springer index of viruses. Springer, New York, pp 1589–1593. https://doi.org/10.1007/978-0-387-95919-1_260 Kang SG, Bong JK, Eun TL, Moo UC (2007) Allexivirus transmitted by eriophyid mites in garlic plants. J Microbiol Biotechnol 17:1833–1840 Karavina C, Gubba A (2017) Iris yellow spot virus in Zimbabwe: incidence, severity and characterization of Alliuminfecting isolates. Crop Prot 94:69–76 Karavina C, Ibaba JD, Gubba A, Pappu HR (2016) First report of Iris yellow spot virus infecting garlic and leek in Zimbabwe. Plant Dis 100:657 Khan I, Sharma A, Kang SS, Kaur G, Dhatt AS (2016) Molecular characterization and identification of Garlic virus D associated with garlic in India. J Plant Pathol 98:607–609 Klukackova J, Navratil M, Vesela M, Havranek P, Safarova D (2004) Occurrence of garlic viruses in the Czech Republic. Acta Fytotechnica et zootech 7:126–128 Klukackova J, Navratil M, Duchoslav M (2007) Natural infection of garlic (Allium sativum L.) by viruses in the Czech Republic. J Plant Dis Prot 114:97–100 Koo BJ, Kang SG, Chang MU (2002) Survey of garlic virus disease and phylogenetic characterization of garlic viruses of the genus Allexivirus isolated in Korea. J Plant Pathol 18:237–243 Lot H, Delecolle B, Boccardo G, Marzachi C, Milne RG (1994) Partial characterization of reovirus-like particles associated with garlic dwarf disease. Plant Pathol 43:537–546 Lot H, Chovelon V, Souche S, Delecolle B (1998) Effects of onion yellow dwarf and leek yellow stripe viruses on symptomatology and yield loss of three French garlic cultivars. Plant Dis 82:1381–1385 Lunello P, Bravo-Almonacid F, Kobayashi K, Helguera M, Nome SF, Mentaberry A, Conci VC (2000) Distribution of Garlic virus A in different garlic production regions of Argentina. J Plant Pathol 82(1):17–21 Mahmoud SYM, Abo-El Maaty SA, El-Borollosy AM, Abdel-Ghaffar MH (2007) Identification of Onion yellow dwarf potyvirus as one of the major viruses infecting garlic in Egypt. Am Eurasian J Agric Environ Sci 2:746–755 Majumder S, Baranwal VK (2009) First report of Garlic common latent virus in garlic from India. Plant Dis 93:106 Majumder S, Baranwal VK (2014) Simultaneous detection of four garlic viruses by multiplex reverse transcription PCR and their distribution in Indian garlic accessions. J Virol Methods 202:34–38 Majumder S, Johari S (2014) First report of Onion yellow dwarf virus and Garlic common latent virus infection in garlic from Nepal. J Plant Pathol 96:S4.117 Majumder S, Baranwal VK, Joshi S (2008) Simultaneous detection of Onion yellow dwarf virus and Shallot latent virus infected leaves and cloves of garlic by duplex RT-PCR. J Plant Pathol 90:369–372 Majumder S, Yadav V, Yakasai MA, Muhammad JY (2016) First report of Garlic common latent virus in garlic from Nigeria. J Plant Pathol 98:684 Majumder S, Yadav V, Yakasai MA, Muhammad JY (2017) First report of Onion yellow dwarf virus in garlic from Nigeria. J Plant Pathol 99:299 Mavric I, Ravnikar M (2005) A carlavirus serologically closely related to Carnation latent virus in Slovenian garlic. Acta Agric Slov 85:343–349

Allium sativum (Garlic)

83

Melo-Filho PA, Nagata T, Dusi AN, Buso JA, Torres AC, Eiras M, Resende RO (2004) Detection of three Allexivirus species infecting garlic in Brazil. Pesq Agrop Brasileira 39:375–340 Mituti T, Marubayashi JM, Moura MF, Krause-Sakate R, Pavan MA (2011) First report of Shallot latent virus in garlic in Brazil. Plant Dis 95:227 Mohamed NA, Young BR (1981) Garlic yellow streak virus, a potyvirus infecting garlic in New Zealand. Ann Appl Biol 97:65–74 Mohammad G, Kawas H, Hall-Safadi B (2007) Survey of garlic viruses in Southern Syria. http://www.damascusu niversity.edu.sy/mag/farm/images/stories/2550.pdf Navratil M, Safarova D, Tkadlecova E, Klukackova J (2006) Molecular characterization of Czech and Chinese Leek yellow stripe virus isolates from garlic. Acta Virol 50:207–210 Oleas A, Arahana V (2016) First report of Leek yellow stripe virus, Shallot latent virus and Onion yellow dwarf virus in garlic from Ecuador. Plant Dis 100:232 Oliveira ML, Hoffmann MIM, Mituti T, Pavan MA, Krause-Sakate R (2014) First report of Garlic virus X in garlic plants in Brazil. Plant Dis 98:1013 Pappu HR, Hellier BC, Dugan FM (2005) First report of Onion yellow dwarf virus, Leek yellow stripe virus, and Garlic common latent virus in garlic in Washington State. Plant Dis 89:205 Parrano L, Afunian M, Pagliaccia D, Douhan G, Vidalakis G (2012) Characterization of viruses associated with garlic plants propagated from different reproductive tissues from Italy and other geographic regions. Phytopathol Mediterr 51:549–565 Perez-Moreno L, Cordova-Rosales Z, Barboza-Corona E, Ramirez-Malagon R, Ramirez-Lua J, Ruis-Castro S, SilvaRosales L (2006) First report of Leek yellow stripe virus in garlic in the state of Guanajuato, Mexico. Plant Dis 90:1458 Perotto MC, Cafrune EE, Conci VC (2010) The effect of additional viral infections on garlic plants initially infected with Allexiviruses. Eur J Plant Pathol 126(4):489–495 Pramesh D, Baranwal VK (2013) Molecular characterization of coat protein gene of Garlic common latent virus isolates from India: an evidence for distinct phylogeny and recombination. Virus Genes 47:189–193 Pramesh D, Islam S, Singh J, Baranwal VK (2012) Serological and PCR detection of Garlic common latent virus associated with garlic accessions in India. Med Plants 4:17–22 Shahraeen N, Lesemann DE, Ghotbi T (2008) Survey for viruses infecting onion, garlic and leek crops in Iran. OEPP/ EPPO Bull 38:131–135 Shiboleth YM, Gal-On A, Koch M, Rabinowitch HD, Salomon R (2001) Molecular characterization of Onion yellow dwarf virus (OYDV) infecting garlic (Allium sativum L.) in Israel. Ann Appl Biol 138(2):187–195 Song SI, Song JT, Chang MU, Lee JS, Choi YD (1997) Identification of one of the major viruses infecting garlic plants, Garlic virus X. Mol Cells 7:705–709 Song SL, Song JT, Kim CH, Lee JS, Choi YD (1998) Molecular characterization of the Garlic virus X genome. J Gen Virol 79:155–159 Stankovic I, Bulajic A, Vucurovic A, Ristic D, Miojevic K, Nikolic D, Krstic B (2012) First report of Tomato spotted wilt virus infecting onion and garlic in Serbia. Plant Dis 96:918 Stefanac Z (1980) Cucumber mosaic virus in garlic. Acta Bot Croat 39:21–26 Stobbs LW, Van Driel L (1999) First report of Onion yellow dwarf virus in Ontario. Plant Dis 83:782 Sumi S, Tsuneyoshi T, Furutani H (1993) Novel shaped viruses isolated from garlic, Allium sativum, possessing a unique genome organization. J Gen Virol 74:1879–1885 Sumi S, Matsumi T, Tsuneyoshi T (1999) Complete nucleotide sequences of garlic viruses A and C, members of the newly ratified genus Allexivirus. Arch Virol 144(9):1819–1826 Sward RJ (1990) Lettuce necrotic yellows rhabdovirus and other viruses infecting garlic. Australas Plant Pathol 19:46–51 Taglienti A, Tiberini A, Manglli A, Rea R, Paoletti S, Taviani P, Tomassoli L (2018) Molecular identification of allexiviruses in a complex mixture of garlic viruses in Latium (central Italy). Eur J Plant Pathol 150:797–801 Testen AL, Mamiro DP, Meulia T, Subedi N, Islam M, Baysal-Gurel F, Miller SA (2014) First report of Leek yellow stripe virus in garlic in Ohio. Plant Dis 98:574 Torres AC, Eiras M, Resende R (2004) Detection of three Allexivirus species infecting garlic in Brazil. Pesq Agrop Brasileira 39:735–740 Torrico AK, Cafrune EE, Conci VC (2010) First report of Shallot latent virus in Garlic in Argentina. Plant Dis 94:915 Tsuneyoshi T, Matsumi T, Deng TC, Sako I, Sumi S (1998) Differentiation of Allium carlaviruses isolated from different parts of the world based on the viral coat protein sequence. Arch Virol 143(6):1093–1107 Van Dijk P (1993) Carlavirus isolates from cultivated Allium species represent three viruses. Neth J Plant Pathol 99:233–257 Voncina D, Curic K, Fabek S, Toth N (2016) First report of Onion yellow dwarf virus, Leek yellow stripe virus and Garlic common latent virus on garlic in Croatia. Plant Dis 100:656

A

84

Allium tuberosum (Garlic chives, Chinese chive)

Vucurovic A, Vucurovic I, Stankovic I, Bulajic A, Nikolic D, Teodorovic S, Krstic B (2015) First report of Garlic common latent virus infecting garlic in Serbia. Plant Dis 99:894 Ward LI, Perez-Egusquiza Z, Fletcher JD, Clover GRG (2009) A survey of viral diseases of allium crops in New Zealand. Australas Plant Pathol 38:533–539 Wei T, Pearson MN, Fletcher JD (2006) Molecular confirmation of New Zealand Garlic yellow streak virus as Leek yellow stripe virus. Australas Plant Pathol 35:341–346 Wylie SJ, Li H, Jones MGK (2012) Phylogenetic analysis of allexiviruses identified on garlic from Australia. Aust Plant Dis Notes 7:23–27 Yamashita K, Sakai J, Hanada K (1995) Leek yellow stripe virus (LYSV) isolated from garlic and its relationship to garlic mosaic (GMV). Ann Phytopathol Soc Jpn 61:273–278 Yamashita K, Sakai J, Hanada K (1996) Characterization of a new virus from garlic (Allium sativum L.), garlic mite-borne virus. Ann Phytopathol Soc Jpn 62:483–489

Allium tuberosum (Garlic chives, Chinese chive) Family: Amaryllidaceae

Spice crop

Cherry leaf roll virus

(CLRV)

Taxonomic position Genus: Nepovirus

Family: Secoviridae

CLRV infection in plants of Allium tuberosum was reported from Finland (AB168098) (von Bargen et al. 2009). The virus is transmitted by nematode vectors in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of CLRV, refer to Prunus avium.

Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

IYSV infection in plants of Allium tuberosum was reported from India and Japan (Fukuda et al. 2007; Gawande et al. 2014). The virus-infected garlic chive plants show straw-colored spindle-like lesions on leaves. The virus is transmitted by onion thrips, Thrips tabaci, in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of IYSV, refer to Iris spp. or Allium cepa.

References Fukuda T, Nakayama K, Honda Y (2007) Necrotic streak disease of Chinese chive caused by Iris yellow spot virus (IYSV). Jpn J Phytopathol 73:311–313 Gawande SJ, Gurav VS, Ingle AA, Gopal J (2014) First report of Iris yellow spot virus infecting Allium tuberosum in India. Plant Dis 98:1161 von Bargen S, Grubits E, Jalkanen R, Buttner C (2009) Cherry leaf roll virus – an emerging virus in Finland? Silva Fenn 43:727–738

Allium spp. (Allium caeruleum, A. christophii, A. giganteum, A. unifolium, and A. “Globemaster”)

85

Allium spp. (Allium caeruleum, A. christophii, A. giganteum, A. unifolium, and A. “Globemaster”) Family: Amaryllidaceae

Allium virus X

Ornamental

(AlVX)

Taxonomic position Genus: Potexvirus

Family: Alphaflexiviridae

Geographical distribution AlVX infection in plants of Allium spp. was reported from the Netherlands (Miglino et al. 2011). Symptoms and host(s) The virus-infected allium plants exhibit diffused yellow stripes or leaf mottling symptoms. Transmission The virus is mechanically sap-transmissible. No vector is known for this virus. Virion properties and genome The virions are flexuous filaments, 470–580 nm in length and 13 nm in diameter. The genome is singlestranded RNA of 7118 nt (FJ670570 = NC_012211) and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type, 18–27 kDa in size (Adams et al. 2004; Miglino et al. 2011).

Garlic virus B

(GarV-B)

Taxonomic position Genus: Allexivirus

Family: Alphaflexiviridae

GarV-B infection in plants of Allium caeruleum was reported from Poland (Paduch-Cichal and Bereda 2017). The virus is transmitted by mite vectors and also by mechanical sap-inoculation. Use of infected vegetative propagative material is another source of virus spread. For more details of GarV-B, refer to Allium sativum.

Leek yellow stripe virus Taxonomic position Genus: Potyvirus

(LYSV)

Family: Potyviridae

LYSV infection in plants of Allium spp. was reported from Japan, the Netherlands, and the UK (Noda and Inouye 1989; Pham et al. 2011; Scrace et al. 2015). The virus-infected allium plants exhibit symptoms of yellow flecking and striping on the leaves. The virus is transmitted by aphid vectors in

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Allium spp. (Allium caeruleum, A. christophii, A. giganteum, A. unifolium, and A. “Globemaster”)

a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of LYSV, refer to Allium ampeloprasum var. porrum.

Narcissus yellow stripe virus Taxonomic position Genus: Potyvirus

(NYSV)

Family: Potyviridae

NYSV infection in plants of Allium carinatum was reported from the USA (Bampi et al. 2018). The virus-infected allium plants show chlorotic streaking symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of NYSV, refer to Narcissus spp.

Onion yellow dwarf virus Taxonomic position Genus: Potyvirus

(OYDV)

Family: Potyviridae

OYDV infection in plants of Allium vineale was reported from the UK (Scrace et al. 2015). The virusinfected allium plants exhibit symptoms of yellow flecking and stripes on the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of OYDV, refer to Allium cepa.

Ornamental onion stripe mosaic virus

(OOSMV)

Taxonomic position OOSMV is a tentative member of the genus Potyvirus and family Potyviridae. Geographical distribution OOSMV infection in plants of the ornamental Allium loratum was reported from the Netherlands and the UK (Pham et al. 2011; Scrace et al. 2015). Symptoms and host(s) The virus-infected allium plants exhibit symptoms of yellow flecking and stripes on the leaves. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are flexuous filaments. The genome is a single molecule of positive-sense single-stranded RNA. A partial polyprotein gene sequence of 1600 nt is available (EU042750) (Wylie et al. 2017).

Allium spp. (Allium caeruleum, A. christophii, A. giganteum, A. unifolium, and A. “Globemaster”)

Pea early-browning virus Taxonomic position Genus: Tobravirus

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(PEBV)

Family: Virgaviridae

PEBV infection in plants of the ornamental Allium karataviense was reported from the UK (Scrace et al. 2015). The virus-infected allium plants exhibit yellow flecking and striping symptoms on the leaves. The virus is transmitted by nematode vectors and also by mechanical sap-inoculation. For more details of PEBV, refer to Pisum sativum.

Shallot latent virus Taxonomic position Genus: Carlavirus

(SLV)

Family: Betaflexiviridae

SLV infection in plants of Allium spp. was reported from the Netherlands (Pham et al. 2011). The virusinfected plants exhibit mosaic and streaking symptoms. The virus is transmitted by aphid vectors in a nonpersistent manner, and also by mechanical sap-inoculation. For more details of SLV, refer to Allium cepa var. aggregatum.

Shallot virus X

(ShVX)

Taxonomic position Genus: Allexivirus

Family: Alphaflexiviridae

ShVX infection in plants of Allium caeruleum was reported from Poland (Bereda and Paduch-Cichal 2016). The virus is transmitted by a mite vector, Aceria tulipae, and also by mechanical sap-inoculation. The use of infected vegetative propagative material is another source of virus spread. For more details of ShVX, refer to Allium cepa var. aggregatum.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in plants of Allium caeruleum was reported from the UK (Scrace et al. 2015). The virusinfected allium plants exhibit yellow flecking and striping symptoms. The virus is transmitted by nematodes of the genera Paratrichodorus and Trichodorus and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

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Allium spp. (Allium bisceptrum, A. canadense, A. validum, A. vineale, A. tricoccum) (Wild onion)

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Allium scorodoprasum was reported from the UK (Scrace et al. 2015). The virus-infected allium plants exhibit chlorosis and yellow striping, with leaf distortion and often necrotic flecking symptoms. The virus is transmitted by aphid vectors Myzus persicae and Brevicoryne brassicae in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TuMV, refer to Brassica rapa.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Bampi D, Reinsel MD, Hammond J (2018) Identification of Narcissus yellow stripe virus and a closely-related Potyvirus isolate in plants of Allium carinatum. Acta Hortic 1193:39–46 Bereda M, Paduch-Cichal E (2016) First report of Shallot virus X in Allium caeruleum in Poland. Plant Dis 100:1958 Miglino R, Druffel KL, van Schadewijk AR, Pappu HR (2011) Molecular characterization of Allium virus X, a new Potexvirus in the family Alphaflexiviridae, infecting ornamental allium. Arch Virol 156:2113–2115 Noda C, Inouye N (1989) Leek yellow stripe virus isolated from an ornamental Allium plant in Japan. Ann Phytopathol Soc Jpn 55:208–215 Paduch-Cichal M, Bereda M (2017) Viruses infecting ornamental allium species in Poland. J Plant Pathol 99(2). https:// doi.org/10.4454/jpp.v99i2.3882 Pham KTK, de Kock MJD, Lemmers MEC, Derks AFLM (2011) Molecular identification of potyviruses infecting bulbous ornamentals by the analysis of coat protein (CP) sequences. Acta Hortic 901:167–172 Scrace J, Denton JO, Clover GRG (2015) Potyviruses and tobraviruses infecting ornamental Allium species in the United Kingdom. New Dis Rep 32:13 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Allium spp. (Allium bisceptrum, A. canadense, A. validum, A. vineale, A. tricoccum) (Wild onion) Family: Amaryllidaceae

Ornamental

Shallot virus X

(ShVX)

Taxonomic position Genus: Allexivirus

Family: Alphaflexiviridae

ShVX infection in plants of Allium vineale was reported from Poland (Paduch-Cichal and Bereda 2017). The virus is transmitted by a mite vector, Aceria tulipae, and also by mechanical sap-inoculation. Use of infected vegetative, propagative material is another source of virus spread. For more details of ShVX, refer to Allium cepa var. aggregatum.

Alocasia spp. (Giant taro)

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Wild onion symptomless virus Taxonomic position Genus: Potyvirus

(WoSV)

Family: Potyviridae

Geographical distribution WoSV infection in plants of Allium spp. was reported from western Turkey (Ohshima et al. 2016). Symptoms and host(s) The virus-infected wild onion plants do not display any external symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner and also by mechanical sapinoculation. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long and 11–13 nm wide. The genome contains single molecule of linear, positive sense single-stranded RNA of 9369 nt in length (LC159494 = NC_030391; LC159495) (Ohshima et al. 2016; Wylie et al. 2017).

References Ohshima K, Korkmaz S, Mitoma S, Nomiyama R, Honda Y (2016) First genome sequence of Wild onion symptomless virus, a novel member of Potyvirus in the Turnip mosaic virus phylogenetic group. Genome Announc 4(4): e00851–e00816. https://doi.org/10.1128/genomeA.00851-16 Paduch-Cichal M, Bereda M (2017) Viruses infecting ornamenal allium species in Poland. J Plant Pathol 99(2). https:// doi.org/10.4454/jpp.v99i2.3882 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Alocasia spp. (Giant taro) Family: Araceae

Dasheen mosaic virus Taxonomic position Genus: Potyvirus

Tuber Crop

(DsMV)

Family: Potyviridae

DsMV infection in plants of Alocasia macrorrhizos was reported from the USA (Elliott et al. 1997). The virus-infected giant taro plants exhibit leaf chlorosis, mosaic, and puckering symptoms (Nelson 2008). The virus is transmitted by aphid vectors such as Myzus persicae, Aphis craccivora, and A. gossypii in a non-persistent manner, and also by mechanical sap-inoculation. For more details of DsMV, refer to Colocasia esculenta.

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Alopecurus myosuroides (Black-grass)

Zantedeschia mild mosaic virus Taxonomic position Genus: Potyvirus

(ZaMMV)

Family: Potyviridae

ZaMMV infection in plants of Alocasia spp. was reported from Australia (Kidanemariam et al. 2016). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. The primary spread of the virus takes place through the use of vegetative planting material. For more details of ZaMMV, refer to Zantedeschia spp.

References Elliott MS, Zettler FW, Brown LG (1997) Dasheen mosaic potyvirus of edible and ornamental aroids. Plant Pathology Circular No. 384, Fla Dept and Consumer Services, July/August 1997 Kidanemariam DB, Abraham AD, Sukal AC, Holton TA, Dale JL, James AP, Harding RM (2016) Complete genome sequence of a novel Zantedeschia mild mosaic virus isolate: the first report from Australia and from Alocasia sp. Arch Virol 161:1079–1082 Nelson SC (2008) Dasheen mosaic of edible and ornamental aroids. Plant Disease CTAHR. Available at http://www.ctahr. hawaii.edu/oc/freepubs/pdf/PD-44.pdf

Alopecurus myosuroides (Black-grass) Family: Poaceae

Weed host

Alopecurus myosuroides varicosavirus 1

(AMVV1)

Taxonomic position AMVV1 is a tentative member of the genus Varicosavirus and family Rhabdoviridae Geographical distribution AMVV1 infection in plants of Alopecurus myosuroides was reported from the UK (Sabbadin et al. 2017). Transmission The virus is transmitted by fungal vector. The virus is not mechanically sap-transmitted. Virion properties and genome The virions are fragile, nonenveloped rods of 13 nm wide and 320–360 nm length. The genome consists of negative-sense, single-stranded RNA. RNA 1 contains 6552 nt (LN713933) and RNA 2 of 4006 nt (LN713934) (Walker et al. 2018).

References Sabbadin F, Glover R, Stafford R, Rozado-Aguirre Z, Boonham N, Adams I, Mumford R, Edwards R (2017) Transcriptome sequencing identifies novel persistent viruses in herbicide resistant wild-grasses. Sci Rep 7:41987. https:// doi.org/10.1038/srep41987 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, ICTV Report Consortium (eds) (2018) ICTV virus taxonomy profile: Rhabdoviridae. J Gen Virol 99:447–448

Alpinia spp. (Flowering ginger)

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Alpinia spp. (Flowering ginger) Family: Zingiberaceae

Ornamental

Alpinia mosaic virus

(AlpMV)

Taxonomic position Genus: Macluravirus

Family: Potyviridae

Geographical distribution AlpMV infection in plants of Alpinia spp. was reported from Taiwan (Chen and Hong 1996). Symptoms and host(s) The virus-infected flowering ginger plants exhibit light green striping symptoms. Transmission The virus is transmitted in a non-persistent manner by the banana aphid, Pentalonia nigronervosa (Chen and Hong 1996). The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped and flexuous filaments 650–675 nm long and 13–16 nm wide. The genome consists of a single molecule of linear positive-sense, single-stranded RNA. A partial polyprotein gene sequence of 1731 nt is available (AF499025) (Liou et al. 2003; Lopez-Moya et al. 2009; Foster 2011; Wylie et al. 2017).

Banana bract mosaic virus Taxonomic position Genus: Potyvirus

(BBrMV)

Family: Potyviridae

BBrMV infection in plants of Alpinia purpurata was reported from Hawaii (Wang et al. 2010; Zhang et al. 2016). The virus-infected flowering ginger plants exhibit symptoms of severe mosaic and stripes on the leaves. Flowers show significant cupping, browning, and reduction in size and shelf life. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BBrMV, refer to Musa spp.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Alpinia galanga was reported from India (Selvarajan et al. 2015). The virus-infected flowering ginger plants exhibit symptoms of spindle-shaped yellow or light green streaks in the interveinal tissues and midribs. The virus is transmitted by aphid vectors in a non-persistent

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Alstroemeria spp.

manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Canna yellow mottle virus Taxonomic position Genus: Badnavirus

(CaYMV)

Family: Caulimoviridae

CaYMV infection in plants of Alpinia purpurata was reported from Hawaii (Zhang et al. 2017). The virus-infected flowering ginger plants exhibit symptoms of yellow mottling and necrosis of leaves, vein streaking, and stunting of plants. The virus spreads through the use of infected planting materials. For more details of CaYMV, refer to Canna spp.

References Chen TH, Hong JL (1996) Zingiberaceous ornamental mosaic- a new disease caused by an aphid-borne potyvirus in Taiwan. Plant Pathol Bull 5:169–175 Foster GD (2011) Macluravirus. Potyviridae. In: The Springer index of viruses. Springer, New York, pp 1421–1424. https://doi.org/10.1007/978-0-387-95919-1_234 Liou RF, Yan HZ, Hong JL (2003) Molecular evidence that aphid-transmitted Alpinia mosaic virus is a tentative member of the genus Macluravirus. Arch Virol 148(6):1211–1218 Lopez-Moya JJ, Valli A, Garcia JA (2009) Potyviridae. In: Iacobellis NS, Collmer A, Hutcheson SW, Mansfield JW, Morris CE, Murillo J, Schaad D NW, Stead E, Surico G, Ullrich M (eds.). Encyclopaedia of life sciences (ELS). Kluwer Academic, Dordrecht, The Netherlands. https://doi.org/10.1002/9780470015902.a0000755.pub2 Selvarajan R, Balasubramanian V, Mandal B (2015) First report of Bean yellow mosaic virus in Alpinia galanga in India. J Plant Pathol 97:541–551 Wang I-C, Sether DM, Melzer MJ, Borth WB, Hu JS (2010) First report of Banana bract mosaic virus in Flowering Ginger in Hawaii. Plant Dis 94:921 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Zhang J, Borth WB, Lin B, Dey KK, Melzer MJ, Shen H, Pu X, Sun D, Hu JS (2016) Deep sequencing of Banana bract mosaic virus from flowering ginger (Alpinia purpurata) and development of an immunocapture RT-LAMP assay. Arch Virol 161:1783–1795 Zhang J, Dey KK, Lin B, Borth WB, Melzer MJ, Sether D, Wang Y, Wang I-C, Shen H, Pu X, Sun D, Hu JS (2017) Characterization of Canna yellow mottle virus in a new host, Alpinia purpurata, in Hawaii. Phytopathology 107:791–799

Alstroemeria spp. Family: Alstroemeriaceae

Ornamental

Alstroemeria mosaic virus

(AlMV)

Synonyms Alstroemeria streak virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

Alstroemeria spp.

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Geographical distribution AlMVoccurs wherever Alstroemeria spp. are grown (Inoue et al. 1992; Wong et al. 1992; Bellardi et al. 1994; van Zaayen et al. 1994; van Zaayen 1995; Bouwen and van der Vlugt 1996; Spence et al. 2000; Wang and Chang 2006). Alstroemeria streak virus has been reported to be a strain of AlMV (van der Vlugt and Bouwen 1999). Symptoms and host(s) The virus-infected alstroemeria plants exhibit streaking of the leaf (chlorosis), which show light green and darker spots parallel to the midrib. Considerable variation in symptom expression is noticed, depending on cultivar, growing conditions, and time of the year. Sometimes a few dark stripes are visible on the flower petals (Phillips and Brunt 1986). Transmission The virus is transmitted by an aphid vector, Myzus persicae, in a non-persistent manner. The virus is also mechanically sap-transmissible to a limited range of indicator plants. The virus is not seed-borne and not transmitted by pollen. Virion properties and genome The virions are non-enveloped, flexuous filaments 780 nm long, and 12 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA. A partial nuclear inclusion protein b/coat protein of 1867 nt (AB158522) and polyprotein gene of 1855 nt have been described (DQ295032) (Wong et al. 1994; Fuji et al. 2004; Revers and Garcia 2015; Wylie et al. 2017).

Alstroemeria necrotic streak virus

(ANSV)

Taxonomic position ANSV is a tentative member of the genus Orthotospovirus and family Tospoviridae. Geographical distribution ANSV infection in plants of Alstroemeria spp. was reported from Colombia (Hassani-Mehraban et al. 2010). Symptoms and host(s) The virus-infected alstroemeria plants exhibit symptoms of necrotic streaks on leaves. Transmission The virus is transmitted by a thrips vector Frankliniella occidentalis in a persistent propagative manner. Virion properties and genome The virions are quasi-spherical, 80–120 nm in diameter, and enveloped by a lipid membrane. The virus contains a single-stranded, tripartite RNA genome, of which the small (S) RNA segment encodes the nucleocapsid (N) protein and a nonstructural NSS protein in an ambisense gene arrangement. A complete nucleocapsid protein (N) mRNA of 1240 nt is available (GQ478668).

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Alstroemeria virus X Taxonomic position Genus: Potexvirus

Alstroemeria spp.

(AlsVX)

Family: Alphaflexiviridae

Geographical distribution AlsVX infection in plants of Alstroemeria spp. was reported from Japan (Fuji et al. 2005). Symptoms and host(s) The virus-infected alstroemeria plants exhibit mosaic symptoms. The virus has a broad host range. Transmission No vector is known for this virus. The virus is mechanically sap-transmissible. Virion properties and genome The virions are flexuous filaments 530 nm in length, and 13 nm in diameter (Fuji et al. 2005). The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 7009 nt (AB206396 = NC_007408) and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type, 18–27 kDa in size (Adams et al. 2004).

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

ArMV infection in plants of Alstroemeria spp. was reported from Italy, Japan, and Lithuania (Bellardi et al. 1986; Inoue et al. 1992; Bellardi et al. 1994; Samuitiene et al. 2008). The virus-infected alstroemeria plants exhibit symptoms of stunting, malformation, and necrotic spots on the leaves. This virus is transmitted by nematode vectors (Xiphinema spp.) in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Broad bean wilt virus 2 Taxonomic position Genus: Fabavirus

(BBWV-2)

Family: Secoviridae

Alstroemeria plants in Japan were occasionally found to be infected with BBWV-2, which induced mosaic in seedling plants (Fuji et al. 2007). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BBWV-2, refer to Vicia faba.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

Alstroemeria spp.

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CMV infection in plants of Alstroemeria spp. was reported from the United Kingdom, India, the Netherlands, and Japan (Spence et al. 2000; Chen 2003; Verma et al. 2005; Fuji et al. 2007). The virusinfected alstroemeria plants exhibit symptoms of interveinal chlorosis, chlorotic mosaic, necrosis, or latent infection. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. The virus was shown to be transmitted through seeds of Alstroemeria pulchella and Alstroemeria hybrids from Italy and the Netherlands (Bellardi and Bertaccini 1997). For more details of CMV, refer to Cucumis sativus.

Freesia mosaic virus Taxonomic position Genus: Potyvirus

(FreMV)

Family: Potyviridae

FreMV infection in plants of Alstroemeria spp. was reported from Italy (Bellardi et al. 1992, 1994; van Zaayen 1995). The virus-infected alstroemeria plants exhibit symptoms of deformed and chlorotic leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of FreMV, refer to Freesia spp.

Grapevine Algerian latent virus Taxonomic position Genus: Tombusvirus

(GALV)

Family: Tombusviridae

GALV infection in plants of Alstroemeria spp. was reported from Japan (Tomitaka et al. 2016). The virus-infected alstroemeria plants exhibit symptoms of necrotic spots on the leaves. There is no known vector for this virus. The virus is mechanically sap-transmissible. The primary spread of the virus takes place through the use of infected planting material. For more details of GALV, refer to Vitis vinifera.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Alstroemeria spp. was reported from Iran (Ghotbi 2013). The virus-infected alstroemeria plants show oval to round chlorotic spots, consisting of many parallel concentric circles, and necrotic spots on the leaves. Stem necrosis and toppling upper portions of plants were not observed (van Zaayen 1995). The virus is transmitted by a thrips vector Frankliniella occidentalis in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of INSV, refer to Impatiens spp.

Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

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Alstroemeria spp.

IYSV infection in plants of Alstroemeria spp. was reported from Japan (Okuda et al. 2003). The virusinfected alstroemeria plants show necrotic streak symptoms (Okuda et al. 2005). The virus is transmitted by thrips vectors in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of IYSV, refer to Iris spp. or Allium cepa.

Lily mottle virus Taxonomic position Genus: Potyvirus

(LMoV)

Family: Potyviridae

LMoV infection in plants of Alstroemeria brasiliensis was reported from the Netherlands (Bouwen and van der Vlugt 1996, 2000a). The virus-infected alstroemeria plants show foliar chlorosis with oval deep green spots and floral color break. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of LMoV, refer to Lilium spp.

Lily symptomless virus

(LSV)

Synonyms Alstroemeria carlavirus

Taxonomic position Genus: Carlavirus

Family: Betaflexiviridae

LSV infection in plants of Alstroemeria spp. was reported from the Netherlands, Italy, and the United Kingdom (Bellardi et al. 1994; Derks et al. 2002; Spence et al. 2000) to cause leaf curling and striping under cool conditions; sequencing of multiple isolates confirmed that the “Alstroemeria carlavirus” is indeed a host-adapted isolate of Lily symptomless virus. The virus is transmitted by an aphid vector Myzus persicae in a non-persistent manner, and also by mechanical sap-inoculation. For more details of LSV, refer to Lilium spp.

Ornithogalum mosaic virus Taxonomic position Genus: Potyvirus

(OrMV)

Family: Potyviridae

OrMV infection in plants of Alstroemeria caryophyllea was reported from the Netherlands (Bouwen and van der Vlugt 1996, 2000b). The virus-infected alstroemeria plants exhibit symptoms of dark green vein banding, necrotic spots, and color breaking in flowers. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of OrMV, refer to Ornithogalum spp.

Alstroemeria spp.

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Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

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Family: Virgaviridae

Infection of alstroemeria with TRV has been reported from the Netherlands (Hakkaart and Versluijs 1985) and the United Kingdom (Phillips and Brunt 1986; Spence et al. 2000). The virus-infected alstroemeria plants exhibit symptoms of yellow flecks, rings, and foliar malformation. Infection is rare and, as transmission is by nematodes, probably limited to field-produced plants. The virus is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tobacco streak virus

(TSV)

Synonyms Alstroemeria ilarvirus Taxonomic position Genus: Ilarvirus

Family: Bromoviridae

Alstroemeria ilarvirus was reported in the UK hybrids of Alstroemeria caryophylla causing faint chlorotic symptoms on leaves. Subsequent studies indicated this virus to be a strain of Tobacco streak virus (TSV) (Brunt and Phillips 1981). The virus is transmitted by the thrips vectors (Frankliniella schultzei, Thrips tabaci), the virus present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible. The virus is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWVoccurs wherever Alstroemeria spp. plants are grown (Bellardi et al. 1994; Takahashi et al. 1998; Tajik et al. 2013). Systemically infected alstroemeria plants exhibit large (5–15 mm), oval to round chlorotic spots on the leaves, and later these parts become necrotic. Necrotic streaks are noticed on leaves, on petioles, and on stems. The virus is transmitted by thrips vectors in a persistent propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting, but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow ring virus

(TYRV)

Taxonomic position TYRV is a tentative member of the genus Orthotospovirus and family Tospoviridae

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Alstroemeria spp.

TYRV infection in plants of Alstroemeria spp. was reported from Iran (Beikzadeh et al. 2011, 2012). The virus-infected alstroemeria plants show necrosis on stems, petioles, flowers, and leaves. The virus is transmitted by a thrips vector Thrips tabaci in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of TYRV, refer to Solanum lycopersicum.

Youcai mosaic virus Taxonomic position Genus: Tobamovirus

(YoMV)

Family: Virgaviridae

YoMV was reported from Alstroemeria spp. in Japan (Fuji et al. 2007). The virus-infected alstroemeria plants exhibit mosaic symptoms. There is no known vector for this virus. The virus is mechanically saptransmissible and also transmissible through contact between plants. For more details of YoMV, refer to Brassica napus.

Zantedeschia mild mosaic virus Taxonomic position Genus: Potyvirus

(ZaMMV)

Family: Potyviridae

ZaMMV infection in plants of Alstroemeria spp. was reported from Italy (Luvisi et al. 2016). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. The primary spread of the virus takes place through the use of vegetative planting material. For more details of ZaMMV, refer to Zantedeschia spp.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Beikzadeh N, Jafarpour B, Rouhani H, Peters D, Hassani Mehraban A (2011) Molecular diagnosis of Tomato yellow ring virus (TYRV) on Alstroemeria in Khorasan Razavi province, Iran. J Plant Prot (Agric Sci Technol) 25:313–314 Beikzadeh N, Bayat H, Jafarpour B, Rohani H, Peters D, Hassani-Mehraban A (2012) Infection of Alstroemeria plants with Tomato yellow ring virus in Iran. J Phytopathol 160:45–47 Bellardi MG, Bertaccini A (1997) Seed transmission of Cucumber mosaic virus in Alstroemeria. Phytopathol Mediterr 36:159–162 Bellardi MG, Canova A, Gelli C (1986) Comparative studies on gladiolus isolates of Arabis mosaic virus (ArMV). Phytopathol Mediterr 25:85–91 Bellardi MG, Vibio M, Bertaccini A (1992) Natural occurrence of Freesia mosaic virus in Alstroemeria sp. Plant Dis 76:643 Bellardi MG, Bertaccini A, Betti L (1994) Survey of viruses infecting Alstroemeria in Italy. Acta Hortic 377:73–80 Bouwen I, van der Vlugt RAA (1996) Identification and characterization of potyviruses of Alstroemeria spp. Acta Hortic 432:72–74 Bouwen I, van der Vlugt RAA (2000a) Natural infection of Alstroemeria brasiliensis with Lily mottle virus. Plant Dis 84:103 Bouwen I, van der Vlugt RAA (2000b) Natural infection of Alstroemeria caryophyllea with Ornithogalum mosaic virus. Plant Dis 84:202 Brunt AA, Phillips S (1981) Report of the glasshouse crops research institute for 1979: 151 Chen YK (2003) Occurence of Cucumber mosaic virus in ornamental plants and perspectives of transgenic control. PhD thesis, Wageningen University, The Netherlands, p 144

Alternanthera spp.

99

Derks AFLM, Lemmers MEC, Konicheva V, Langeveld SA (2002) Lily symptomless virus in Alstroemeria: Identification and transmission to lily. Acta Hortic 568:247–251 Fuji S, Terami F, Furuya H, Naito H, Fukumoto F (2004) Nucleotide sequence of the coat protein genes of Alstroemeria mosaic virus and Amazon lily mosaic virus, a tentative species of genus Potyvirus. Arch Virol 149:1843–1849 Fuji S, Shinoda K, Ikeda M, Furuya H, Naito H, Fukumoto F (2005) Complete nucleotide sequence of the new Potexvirus “Alstroemeria virus X”. Arch Virol 150:2377–2385 Fuji SI, Mochizuki N, Fujinaga M, Ikeda M, Shinoda K, Uematsu S, Furuya H, Naito H, Fukumoto F (2007) Incidence of viruses in Alstroemeria plants cultivated in Japan and characterization of Broad bean wilt virus-2, Cucumber mosaic virus and Youcai mosaic virus. J Gen Plant Pathol 73:216–221 Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iranian J Plant Pathol 49:41 Hakkaart FA, Versluijs JMA (1985) Viruses of Alstroemeria and preliminary results of meristem culture. Act Hortic 164:71–75 Hassani-Mehraban A, Botermans M, Verhoeven JT, Meekes E, Saaijer J, Peters D, Goldbach R, Kormelink R (2010) A distinct Tospovirus causing necrotic streak on Alstroemeria sp. in Colombia. Arch Virol 155:423–428 Inoue N, Maeda T, Huttinga H (1992) Two viruses, Arabis mosaic virus and Alstroemeria mosaic virus, isolated from alstroemeria. Ann Phytopathol Soc Jpn 58:135 Luvisi A, Rizzo D, Stefani L, Panattoni A, Materazzi A (2016) Occurrence of viruses in Calla and Peruvian lily in Tuscan nurseries and evidence of new viral records in Italy. Adv Hortic Sci 30:53–56 Okuda M, Hanada K, Uematsu S, Fukumoto F, Mihara F, Ebihara T, Iwanami T (2003) Characterization of Iris yellow spot virus isolates causing yellow necrosis on Alstroemeria species. Jpn J Phytopathol 69:23 Okuda M, Hanada K, Uematsu S, Fukumoto F, Mihara T, Ebihara Y, Iwanami T (2005) Necrotic streaks of alstroemeria (Alstroemeria spp.) caused by Iris yellow spot virus (IYSV). Jpn J Phytopathol 71:119–122 Phillips S, Brunt AA (1986) Four viruses of Alstroemeria in Britain. Acta Hortic 177:227–283 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Samuitiene M, Navalinskiene M, Jackeviciene E (2008) Arabis mosaic virus on ornamental plants. Biologia 54:264–268 Spence NJ, Mills PR, Barbara DJ (2000) A survey of virus of Alstroemeria in the UK and characterization of Carlavirus infecting Alstroemeria. Eur J Plant Pathol 106:843–847 Tajik S, Maleki M, Ghotbi T (2013) First report of Tomato spotted wilt virus (TSWV) in Alstroemeria from Iran. Plant Dis. https://doi.org/10.1094/PDIS-09-12-0900-PDN Takahashi Y, Oda K, Takahashi K, Kawano T (1998) Tomato spotted wilt virus (TSWV) isolated from Alstroemeria hybrid showing necrotic lesion (in Japanese). Annu Report Kanto Plant Prot Soc 45:105–108 Tomitaka Y, Usugi T, Fukami M, Tsuda S (2016) First report of Grapevine Algerian latent virus infection on Alstroemeria spp. in Japan. Plant Dis 100:1251 van der Vlugt RAA, Bouwen I (1999) Alstroemeria streak virus is an isolate of Alstroemeria mosaic potyvirus. Phytomedizin Mitteilungen der Deutschen Phytomedizinischen Gesellschaft e.V. Sonderheft 1:31 van Zaayen A (1995) Alstroemeria. In: Loebenstein G, Lawson RH, Brunt AA (eds) Virus and virus like disease of bulb and flower crops. Wiley, Chichester, pp 237–249 van Zaayen A, de Blank CM, Bouwen I (1994) Differentiation between two potyviruses in Alstroemeria. Eur J Plant Pathol 100:85–90 Verma N, Singh AK, Singh L, Raikhy G, Kulshreshta S, Singh MK, Hallan V, Ram R, Zaidi AA (2005) Cucumber mosaic virus (CMV) infecting alstroemeria hybrids in India. Australas Plant Pathol 39:119–120 Wang C-Y, Chang Y-C (2006) First identification of Alstroemeria mosaic virus in Taiwan. Plant Pathol 55:566 Wong SM, Reiser RA, Horst RK (1992) A new virus in Alstroemeria in the USA. Phytopathology 82:722 Wong SM, Chng CG, Reiser RA, Horst RK (1994) Further characterization of a potyvirus in alstroemeria endowment series. Acta Hortic 377:63–71 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Alternanthera spp. Family: Amaranthaceae

Ornamental

Alligatorweed stunting virus Taxonomic position Genus: Unassigned

(AWSV)

Family: Closteroviridae

A

100

Alternanthera spp.

Geographical distribution AWSV infection in plants of Alternanthera philoxeroides was first reported from Florida, USA, by Hill and Zettler (1973). The virus spreads in the North American region in the USA (Florida). Symptoms and host(s) The virus-infected alternanthera plants exhibit leaf reddening and malformation and severe stunting symptoms. Transmission The virus is transmitted by mechanical sap-inoculation. Virion properties and genome The virions are filamentous and usually flexuous with a clear modal length of 1717 nm. The genome is a monopartite, linear, positive-sense, single-stranded RNA.

Alternanthera mild mosaic virus Taxonomic position Genus: Potyvirus

(AltMMV)

Family: Potyviridae

Geographical distribution AltMMV infection in plants of Alternanthera tenella (syn. A. ficoidea) was reported from Brazil (Almeida et al. 2007). Symptoms and host(s) The virus-infected alternanthera plants exhibit mild mosaic symptoms. The virus has a very narrow host range. Transmission The virus is transmitted by an aphid vector Myzus persicae in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments 756 nm long, and 12 nm wide. The genome consists of a single molecule of linear, positive-sense, single-stranded RNA. A partial polyprotein gene sequence of 1772 nt is available (EF442668) (Almeida et al. 2007; Revers and Garcia 2015; Wylie et al. 2017).

Alternanthera mosaic virus Taxonomic position Genus: Potexvirus

(AltMV)

Family: Alphaflexiviridae

Geographical distribution AltMV was first identified in 1999 in Queensland, Australia (Geering and Thomas 1999). The virus was found in Alternanthera pungens (Amaranthaceae), a weed found in both the Southern USA and Australia.

Alternanthera spp.

101

Symptoms and host(s) The virus-infected alternanthera plants exhibit mottle or mosaic symptoms, but necrosis may be induced in some hosts, especially at low temperatures. The virus naturally infects many ornamental hosts including Phlox stolonifera, Portulaca grandiflora, Scutellaria longifolia, Crossandra infundibuliformis, Torenia spp., and Angelonia spp. Since then the virus has been found to infect species in at least 20 taxonomically diverse families, including mainly herbaceous ornamentals but also the woody shrub Nandina domestica (Hammond et al. 2017). To date, there are no reports of natural infections in agronomic crops, but the broad experimental host range indicates that AltMV is able to infect several important vegetable crops (Hammond et al. 2017). Transmission No vector is reported for this virus. The virus is mechanically sap-transmissible and systemically infects many test plants, including tomato, faba bean, sunflower, zinnia, watermelon, cucumbers, and cowpea. Potexviruses are readily sap-transmissible and are easily spread by plant contact, vegetative propagation, and contaminated tools and hands. Some have been shown to retain infectivity in plant sap for several months at room temperature. The virus is not transmitted by seed or pollen. Virion properties and genome The virions are flexuous filaments particles, approximately 536 nm long and 13 nm in diameter. The genome is a single molecule of linear, positive-sense, single-stranded RNA of 6607 nt (AY863024 = NC_007731) and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type, 18–27 kDa in size (Adams et al. 2004).

Alternanthera yellow vein virus Taxonomic position Genus: Begomovirus

(AlYVV)

Family: Geminiviridae

Geographical distribution AlYVV infection in plants of Alternanthera philoxeroides was reported from China (Guo and Zhou 2005). Symptoms and host(s) The virus-infected alternanthera plants exhibit yellow vein symptoms. However, care needs to be taken in interpreting symptoms since these are usually determined by the betasatellite with which the virus associates. AlYVV has been shown to be capable of associating with multiple different betasatellites. The virus has been isolated from Alternanthera philoxeroides, Eclipta prostrata, Ludwigia hyssopifolia, and Zinnia elegans. Transmission The transmission of AlYVV has not been investigated. It is likely that in common with other begomoviruses, virus will be transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Similarly, it is likely that the virus will prove not to be either mechanically or seed-transmitted.

A

102

Alternanthera spp.

Virion properties and genome The structure of the virions of AlYVV has not been investigated. In common with all geminiviruses, the virions of AlYVV are likely geminate (twinned icosahedra). The characterized genomes of AlYVV are ~2745 nt (AJ965540 = NC_007211). The characterized genomes of AlYVV encode the six genes typically encoded by monopartite begomoviruses. The expression and function of the genes have not been investigated. Alternanthera yellow vein betasatellite DNA molecule is associated with the virus, and consists of 1344 nt (DQ641716) (Briddon 2001; Ha et al. 2008; Zhou 2013; Brown et al. 2015; Zerbini et al. 2017).

Iresine viroid 1

(IrVd-1)

Taxonomic position Genus: Pospiviroid

Family: Pospiviroidae

IrVd-1 infection in plants of Alternanthera sessilis was reported from India (Singh et al. 2006). The viroid is mechanically sap-transmissible. For more details of IrVd-1, refer to Iresine spp.

Tomato chlorotic dwarf viroid Taxonomic position Genus: Pospiviroid

(TCDVd)

Family: Pospiviroidae

TCDVd infection in plants of Alternanthera sessilis was reported from India (Singh et al. 2006). The viroid is mechanically sap-transmissible. For more details of TCDVd, refer to Solanum lycopersicum.

Tomato leaf curl Kerala virus

(ToLCKeV)

Synonyms Alternanthera leaf curl virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

ToLCKeV infection in plants of Alternanthera sessilis was reported from India (Marwal et al. 2013). The virus-infected alternanthera plants exhibit leaf curl and chlorosis symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner. For more details of ToLCKeV, refer to Solanum lycopersicum.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Almeida AM, Fukushigue CY, Sartori F, Binneck E, Marin SR, Inoue-Nagata AK, Chagas CM, Souto ER, Mituti T (2007) Natural infection of Alternanthera tenella (Amaranthaceae) by a new potyvirus. Arch Virol 152:2095–2099

Althaea officinalis (Marshmallow)

103

Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Geering AD, Thomas JE (1999) Characterisation of a virus from Australia that is closely related to Papaya mosaic potexvirus. Arch Virol 144:577–592 Guo X, Zhou X (2005) Molecular characterization of Alternanthera yellow vein virus: a new Begomovirus species infecting Alternanthera philoxeroides. J Phytopathol 153:694–696 Ha C, Coombs S, Revill P, Harding R, Vu M, Dale J (2008) Molecular characterization of begomoviruses and DNA satellites from Vietnam: additional evidence that the New World geminiviruses were present in the Old World prior to continental separation. J Gen Virol 89:312–326 Hammond J, Kim I-H, Lim H-S (2017) Alternanthera mosaic virus – an alternative ‘model’ Potexvirus of broad relevance. Kor J Agric Sci 44:145–180 Hill HR, Zettler FW (1973) A virus-like stunting disease of alligatorweed from Florida. Phytopathology 63:443 Marwal A, Sahu AK, Prajapat R, Choudhary DK, Gaur RK (2013) Molecular and recombinational characterization of Begomovirus infecting an ornamental plant Alternanthera sessilis: a new host of Tomato leaf curl kerala virus reported in India. Sci Int 1:51–56 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Singh RP, Dilworth AD, Baranwal VK, Gupta KN (2006) Detection of Citrus exocortis viroid, Iresine viroid, and Tomato chlorotic dwarf viroid in new ornamental host plants in India. Plant Dis 90:1457 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zhou X (2013) Advances in understanding begomovirus satellites. Annu Rev Phytopathol 51:357–381

Althaea officinalis (Marshmallow) Family: Malvaceae

Medicinal

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Althaea officinalis was reported from Bulgaria (Dikova 2011). The virusinfected marshmallow plants exhibit chlorotic/necrotic spotting on leaves. The virus is transmitted by thrips vectors in a persistent, propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting, but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Dikova B (2011) Tomato spotted wilt virus on some medicinal and essential oil-bearing plants in Bulgaria. Bulgarian J Agr Sci 17:306–313

A

104

Althaea rosea (Hollyhock)

Althaea rosea (Hollyhock) Synonyms Alcea rosea Family: Malvaceae

Ornamental

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

ArMV infection in plants of Althaea rosea was reported from Iran (Ghotbi and Shahraeen 2012). The virus is transmitted by nematode vectors (Xiphinema spp.) in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Cotton leaf curl Gezira virus

(CLCuGeV)

Synonyms Althea rosea enation virus; Hollyhock leaf crumple virus (HoLCrV) Taxonomic position Genus: Begomovirus

Family: Geminiviridae

HoLCrV infection in plants of Althaea rosea was reported from Egypt (Abdel-Salam et al. 1998; Bigarre et al. 2001). The virus-infected hollyhock plants show crumpling and cupping of leaf blades, thickening of small leaf veins, and epinasty. The virus is transmitted by the whitefly vector Bemisia argentifolii or Bemisia tabaci in a circulative non-propagative manner. For more details of CLCuGeV, refer to Gossypium spp.

Hollyhock leaf curl virus Taxonomic position Genus: Begomovirus

(HoLCV)

Family: Geminiviridae

Geographical distribution HoLCV infection in plants of Alcea rosea was reported from Pakistan (Zia Ur Rehman and Haider, unpublished - FR772082). Symptoms and host(s) The virus-infected hollyhock plants exhibit leaf curl symptoms. The virus has also been isolated from Andrographis paniculata (with leaf curling, vein-clearing, chlorosis, and stunting) and Eclipta prostrata with vein yellowing symptoms.

Althaea rosea (Hollyhock)

105

Transmission The transmission of HoLCV has not been investigated. It is likely that, in common with other begomoviruses, the virus is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of HoLCV has not been investigated. In common with all geminiviruses, the virions of HoLCV are likely geminate (twinned quasi-icosahedra). HoLCV is a typical Old World monopartite begomovirus. The genome of HoLCV consists of a single circular molecule of single-stranded DNA of ~2748 nt (FR772082; GQ478343, KC476655). The characterized genomes of HoLCV encode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated. At least one isolate of HoLCV was reported to be associated with a betasatellite (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Hollyhock yellow vein mosaic virus Taxonomic position Genus: Begomovirus

(HoYVMV)

Family: Geminiviridae

Geographical distribution HoYVMV was first detected in plants of Althaea rosea in Pakistan (FR772081; LM645009) and later in India (Snehi et al. 2013; Srivastava et al. 2014). Symptoms and host(s) The virus-infected hollyhock plants exhibit yellow veinal mosaic symptoms. The plants in India were found to be infected with HoYVMV together with Ludwigia leaf distortion betasatellite (Srivastava et al. 2014). Transmission The transmission of HoYVMV has not been investigated. It is likely that, in common with other begomoviruses, HoYVMV is transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Virion properties and genome The structure of the virions of HoYVMV has not been investigated. In common with all geminiviruses, the virions of HoYVMV are likely geminate (twinned quasi-icosahedra). The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2741 nt (LM645009; FR772081 = NC_016649; JQ911766) (Briddon 2001; Srivastava et. al. 2014; Brown et al. 2015; Zerbini et al. 2017).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(INSV)

A

106

Althaea rosea (Hollyhock)

INSV infection in plants of Althaea spp. was reported from Iran (Ghotbi and Shahraeen 2012; Ghotbi 2013). The virus-infected hollyhock plants exhibit symptoms, including necrotic leaf spot, chlorosis, and stunting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Malva vein clearing virus Taxonomic position Genus: Potyvirus

(MVCV)

Family: Potyviridae

MVCV infection in plants of Alcea rosea was reported from Germany (Menzel et al. 2010). The virusinfected hollyhock plants exhibit strong vein-clearing and vein-yellowing symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of MVCV, refer to Malva sylvestris.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV was detected in plants of Althaea spp. in Iran (Ghotbi and Shahraeen 2009). The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Althaea spp. was reported from Iran (Ghotbi 2006). The virus-infected hollyhock plants exhibit symptoms of mosaic and leaf necrotic lesions. The virus is transmitted by thrips vectors; the virus present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible. The virus is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

Plants of Althaea spp. were found infected with ToRSV in Iran (Ghotbi and Shahraeen 2012). The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation, and by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Althaea rosea (Hollyhock)

107

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

Plants of Althaea spp. were found infected with TSWV in Iran (Parrella et al. 2003; Ghotbi and Shahraeen 2012). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting, but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow ring virus

(TYRV)

Taxonomic position TYRV is a tentative member of the genus Orthotospovirus and family Tospoviridae Althaea spp. plants were reported to be infected with TYRV in Iran (Ghotbi and Shahraeen 2012). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sapinoculation. For more details of TYRV, refer to Solanum lycopersicum.

Zucchini yellow mosaic virus Taxonomic position Genus: Potyvirus

(ZYMV)

Family: Potyviridae

ZYMV infection in plants of Alcea rosea was reported from Korea (Choi et al. 2002, 2007). The virusinfected hollyhock plants exhibit mild mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ZYMV, refer to Cucurbita pepo.

References Abdel-Salam AM, El-Shazly MA, Thouvenel JC (1998) Biological, biochemical and serological studies on Hollyhock leaf crumple cirus (HLCrV): a newly discovered whitefly transmitted geminivirus. Arab J Biotechnol 1:41–58 Bigarre L, Chazly M, Salah M, Ibrahim M, Padidam M, Nicole M, Peterschmitt M, Fauquet C, Thouvenel JC (2001) Characterization of a new begomovirus from Egypt infecting hollyhock (Althea rosea). Eur J Plant Pathol 107:701–711 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Choi SK, Yoon JY, Ryu KH, Choi JK, Mok W (2002) First report of Zucchini yellow mosaic virus on Hollyhock (Althaea rosea). Plant Pathol J 18:121–125 Choi SK, Yoon JY, Sohn SH (2007) Analysis of the complete genome sequence of Zucchini yellow mosaic virus strain A isolated from hollyhock. Plant Pathol J 23:245–250 Ghotbi T (2006) First report on incidence of Tobacco streak virus (TSV) on ornamental plants in Iran. Iranian J Plant Pathol 42:159–160

A

108

Alysicarpus spp. (Alyce clover)

Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iran J Plant Path 49:41 Ghotbi T, Shahraeen N (2009) Natural incidence and infectivity level of three nepoviruses in ornamental crops in Iran. J Plant Breed Crop Sci 1:39–44 Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses affecting propagated ornamentals in Northern Iran. Int Res J Microbiol 3:373–381 Menzel W, Winter S, Richert-Poggeler KR (2010) First report of Malva vein clearing virus naturally occurring in hollyhock in Germany. Plant Dis 94:276 Parrella G, Gognalons P, Gebre-Selassie K, Vovlas C, Marchoux G (2003) An update of the host range of Tomato spotted wilt virus. J Plant Pathol 85:227–264 Snehi SK, Srivastava A, Kumar S, Raj SK (2013) Molecular detection and identification of begomoviruses affecting important ornamental plants in India. Indian J Virol 24:142 Srivastava A, Kumar S, Raj SK, Pande SS (2014) Association of a distinct strain of Hollyhock yellow vein mosaic virus and Ludwigia leaf distortion betasatellite with yellow vein mosaic disease of hollyhock (Alcea rosea) in India. Arch Virol 159:2711–2715 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Alysicarpus spp. (Alyce clover) Family: Fabaceae

Fodder crop

Bean common mosaic virus

(BCMV)

Synonyms Blackeye cowpea mosaic virus (BlCMV) Taxonomic position Genus: Potyvirus

Family: Potyviridae

BlCMV-infected plants of Alysicarpus vaginalis were reported from Florida (USA) (Zhao et al. 1991a, b). The virus-infected Alyce clover plants exhibit mosaic, stunting, and leaf distortion symptoms. The virus is transmitted by an aphid vector Myzus persicae in a non-persistent manner, and also by mechanical sapinoculation. For more details of BCMV, refer to Phaseolus vulgaris.

Clover yellow vein virus Taxonomic position Genus: Potyvirus

(ClYVV)

Family: Potyviridae

ClYVV infection in plants of Alysicarpus vaginalis was reported from Georgia (USA) (Miller et al. 1987). The virus-infected Alyce clover plants exhibit yellow vein symptoms. The virus is transmitted by several aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ClYVV, refer to Trifolium spp.

Amaranthus spp.

109

Peanut mottle virus Taxonomic position Genus: Potyvirus

(PeMoV)

Family: Potyviridae

PeMoV infection in plants of Alysicarpus vaginalis was reported from Georgia (USA) (Miller et al. 1987). The virus-infected Alyce clover plants exhibit mottling symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PeMoV, refer to Arachis hypogaea.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Alysicarpus muelleri was reported from Australia (Sharman et al. 2015). The virus-infected plants exhibit symptoms of chlorotic line patterns and mottling symptoms. The virus is transmitted by thrips vectors, the virus present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

References Miller JD, Demski JW, Baltensperger DD, Wells HD (1987) First report of Peanut mottle virus and Clover yellow vein virus on alyce-clover in Georgia. Plant Dis 71:192 Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct Tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207 Zhao GS, Baltensperger DD, Hiebert E, Purcifull DE, Edwardson JR (1991a) Purification, serology, and in vitro translation of an alyce-clover isolate of Blackeye cowpea mosaic virus. Plant Dis 75:254–257 Zhao GS, Baltensperger DD, Purcifull DE, Christie RG, Hiebert E, Edwardson JR (1991b) Host range, cytology and transmission of an alyce-clover isolate of Black eye cowpea mosaic virus. Plant Dis 75:251–253

Amaranthus spp. Family: Amaranthaceae

Ageratum enation virus Taxonomic position Genus: Begomovirus

Leafy vegetable

(AEV)

Family: Geminiviridae

AEV infection in plants of Amaranthus spp. was reported from India (Raj et al. 2008; Srivastava et al. 2013, 2014). The virus-infected amaranthus plants exhibit symptoms of upward leaf curling, vein enation on the lower side of the leaves, and shortening of internodes. The virus is transmitted by the

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Amaranthus spp.

whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. The virus is not transmitted by mechanical sap-inoculation. For more details of AEV, refer to Ageratum spp.

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

(AMV)

Family: Bromoviridae

AMV infection in plants of Amaranthus albus was reported from the USA (Kaiser and Hannan 1983). The virus-infected amaranthus plants exhibit symptoms of stunt, and the foliage shows yellow mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is also seed-transmitted in A. albus to the extent of 1.9–15.5% (Kaiser and Hannan 1983). For more details of AMV, refer to Medicago sativa.

Amaranthus leaf mottle virus Taxonomic position Genus: Potyvirus

(AmLMV)

Family: Potyviridae

Geographical distribution AmLMV infection was first reported in Amaranthus deflexus plants from Turin, Italy, by Lovisolo and Lisa (1976). The virus spreads in the Eurasian regions: Italy, Morocco, and Spain (Lovisolo and Lisa 1979; Segundo et al. 2007). Symptoms and host(s) The virus-infected amaranthus plants exhibit symptoms of leaf mottling, blistered mosaic, and growth reduction. Transmission The virus is transmitted by aphid vectors, Macrosiphum euphorbiae and Myzus persicae, in a nonpersistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments with a clear modal length of 780 nm and 12 nm wide. The genome consists of a single molecule of linear, positive-sense, single-stranded RNA. A partial polyprotein gene of 1777 nt is available (AJ580095) (Segundo et al. 2007; Revers and Garcia 2015; Wylie et al. 2017).

Amaranthus mosaic virus Taxonomic position The virus is a tentative member of the genus Potyvirus and family Potyviridae.

Amaranthus spp.

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Geographical distribution This virus infection in plants of Amaranthus spp. was reported from India and Nigeria (Phatak 1965; Govindasamy et al. 1967; Ramakrishnan et al. 1971; Taiwo 1988; Awasthi and Khare 2006). Symptoms and host(s) The virus-infected amaranthus plants exhibit severe mosaic, mottling, and curling leaves with stunting. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically saptransmissible. Virion properties and genome The virions are flexuous filaments measuring 700–760 nm in length and 14 nm wide. The genome is a single molecule of positive-sense, single-stranded RNA.

Beet curly top virus Taxonomic position Genus: Curtovirus

(BCTV)

Family: Geminiviridae

BCTV infection in plants of Amaranthus albus was reported from California (USA) (Creamer et al. 1996). The virus is transmitted by the leafhopper vector, Circulifer tenellus, in a persistent (circulative, non-propagative) manner. The virus is not transmitted by mechanical inoculation. For more details of BCTV, refer to Beta vulgaris.

Capsicum chlorosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(CaCV)

Family: Tospoviridae

CaCV infection in plants of Amaranthus spp. was reported from India (Sharma and Kulshrestha 2014). The virus-infected amaranthus plants exhibit symptoms of curling and chlorosis of its leaves. The virus is transmitted by thrips vectors in a persistent propagative manner, and also by mechanical sapinoculation. For more details of CaCV, refer to Capsicum annuum.

Chilli leaf curl virus Taxonomic position Genus: Begomovirus

(ChiLCV)

Family: Geminiviridae

ChiLCV infection in plants of Amaranthus spp. was reported from India (George et al. 2014). The virusinfected amaranthus plants exhibit symptoms of leaf curling, leaf distortion, leaf crinkling, and yellow leaf margins. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, nonpropagative manner. For more details of ChiLCV, refer to Capsicum annuum.

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Amaranthus spp.

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

CGMMV infection in plants of Amaranthus spp. was reported from Israel (Shargil et al. 2017). The virus is transmissible by mechanical sap-inoculation, and also by contact between plants. For more details of CGMMV, refer to Cucumis sativus.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Amaranthus spp. was reported from India and Japan (Sharma and Chowfla 1986; Vicchi and Bellardi 1988; Ohashi and Kamiunten 1994; Raj et al. 1997, 2000). The virus-infected amaranthus plants show leaf crinkling, leaf deformation, severe mosaic, and stunting that leads to seed yield losses. The virus is mechanically sap-transmissible to a number of host plants, and also several aphid species transmit this virus in a non-persistent manner. For more details of CMV, refer to Cucumis sativus.

Cucurbit yellow stunting disorder virus Taxonomic position Genus: Crinivirus

(CYSDV)

Family: Closteroviridae

CYSDV infection in plants of Amaranthus lividus was reported from Florida (Webster et al. 2011). The virus-infected amaranthus plants exhibit yellowing of leaves and stunting of plants. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a semi-persistent manner. The virus is not mechanically sap-transmissible. For more details of CYSDV, refer to Cucumis melo.

Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

IYSV infection in plants of Amaranthus spp. was reported from Zimbabwe (Karavina and Gubba 2017). The virus is transmitted by onion thrips, Thrips tabaci, in a persistent, propagative manner, and also by mechanical sap-inoculation. For more details of IYSV, refer to Iris spp. or Allium cepa.

Papaya leaf curl virus Taxonomic position Genus: Begomovirus

(PaLCuV)

Family: Geminiviridae

Amaranthus spp.

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PaLCuV infection in plants of Amaranthus cruentus was reported from India (Srivastava et al. 2015). The virus-infected plants exhibit leaf curling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. For more details of PaLCuV, refer to Carica papaya.

Pepino mosaic virus Taxonomic position Genus: Potexvirus

(PepMV)

Family: Alphaflexiviridae

PepMV infection in plants of Amaranthus spp. was reported from Spain (Jorda et al. 2001). The virus is mechanically sap-transmissible. For more details of PepMV, refer to Solanum muricatum.

Potato virus S Taxonomic position Genus: Carlavirus

(PVS)

Family: Betaflexiviridae

PVS infection in plants of Amaranthus hybridus was reported from Iran (Hosseini and Salari 2017). The virus is transmitted by aphid vectors in a non-persistent manner, and also through mechanical sapinoculation. For more details of PVS, refer to Solanum tuberosum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Amaranthus cruentus was reported from Iran and Australia (Ghotbi 2006; Sharman et al. 2015). The virus-infected amaranthus plants exhibit symptoms of vein necrotic lesions and leaf deformation. The virus is transmitted by thrips vectors, the virus present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible. The virus is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Amaranthus retroflexus was reported from South Africa, Georgia (USA) and Italy (Mullis et al. 2009; Grieco et al. 2000; Kisten et al. 2016). The virus is transmitted by thrips vectors in a persistent, propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting, but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

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Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Amaranthus retroflexus was reported from South Florida and Cyprus (Ying and Davis 2000; Papayiannis et al. 2011). The virus-infected amaranthus plants exhibit symptoms of curling, distortion, and chlorosis of leaves, stunting of the plant, and a high rate of flower abscission. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, nonpropagative manner. The virus is transmissible by grafting, but not transmissible by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

References Awasthi LP, Khare MN (2006) Transmission of Amaranthus mosaic virus by bugs from India. Plant Dis Res 21:172–174 Creamer R, Lugue-Williams M, Howo M (1996) Epidemiology and incidence of Beet curly top geminivirus in naturally infected weed hosts. Plant Dis 80:533–535 George B, Vinoth Kumar R, Chakraborty S (2014) Molecular characterization of Chilli leaf curl virus and satellite molecules associated with leaf curl disease of Amaranthus spp. Virus Genes 48:397–401 Ghotbi T (2006) First report on incidence of Tobacco streak virus (TSV) on ornamental plants in Iran. Iranian J Plant Pathol 42:159–160 Govindasamy CV, Marappan V, Kandaswamy TK, Padmanabham C (1967) Mosaic disease of Amaranthus in Coimbatore (Madras State). Madras Agric J 54:35–36 Grieco PD, Conte D, Munno I, Nuzzaci M, de Stradis A (2000) Tomato spotted wilt virus (TSWV) on weeds and wild plants in Metapontino, Basilicata. Informatore Fitopatologico 50:43–46 Hosseini SA, Salari K (2017) Detection and molecular characterisation of Potato virus S of weed reservoirs in Iran. Arch Phytopathol Plant Protect 50:828–838 Jorda C, Lazaro Perez A, Martinez-Culebras P, Lacasa A (2001) First report of Pepino mosaic virus on natural hosts. Plant Dis 85:1292 Kaiser WJ, Hannan RM (1983) Additional hosts of Alfalfa mosaic virus and its seed transmission in tumble pigweed and bean. Plant Dis 67:1354–1357 Karavina C, Gubba A (2017) Amaranthus sp. and Eleusine indica are natural hosts of Iris yellow spot virus in Zimbabwe. Plant Dis 101:262 Kisten L, Moodley V, Gubba A, Mafongoya PL (2016) First detection of Tomato spotted wilt virus (TSWV) on Amaranthus thunbergii in South Africa. Plant Dis 100:2176 Lovisolo O, Lisa V (1976) Characterization of a virus isolated from Amaranthus deflexus, serologically related to Bean yellow mosaic virus. Agric Scient 39:553–559 Lovisolo O, Lisa V (1979) Studies on Amaranthus leaf mottle virus (ALMV) in the Mediterranean region. Phytopathol Mediterr 18:89 Mullis SW, Nischwitz C (2005) Tospovirus detection. In: Tospoviruses in Solanaceae and other crops in the coastal plain of Georgia. University of Georgia, College of Agricultural and Environmental Sciences, Bulletin 1354 Mullis S, Bertrand PF, Brown SL, Csinos A, Diaz-Perez JC, Gitaitis R, Hickman LL, Johnson A, LaHue SS, Martinez N, McPherson RM, Nischwitz C, Reay-Jones F, Riley D, Sherwood J, Stevenson K, Wells L (2009) Tospoviruses in Solanaceae and other crops in the coastal plain of Georgia. University of Georgia, College of Agricultural and Environmental Sciences, Bulletin 1354, pp 51 Ohashi M, Kamiunten H (1994) Mosaic disease of grain amaranth (Amaranthus hypochondriacus L.) caused by Cucumber mosaic virus (CMV). Jpn J Phytopathol 60:119–121 Papayiannis L, Katis NI, Idris AM, Brown JK (2011) Identification of weed hosts of Tomato yellow leaf curl virus in Cyprus. Plant Dis 95:120–125 Phatak HS (1965) Mosaic disease of amaranthus – a new record for India. Curr Sci 34:645–646 Raj SK, Aminuddin SBP, Pal M (1997) Characterization of a Cucumber mosaic virus isolate causing leaf crinkle and severe mosaic of Amaranthus in India. Can J Plant Pathol 19:97–100 Raj SK, Aminuddin RM, Pandey NP (2000) Screening and field evaluation for natural occurrence of Cucumber mosaic virus in Amaranths and Celosia. Indian J Plant Pathol 18:14–18 Raj SK, Snehi SK, Khan MS, Chandra P, Pandey RM (2008) First molecular identification of a Begomovirus associated with yellow vein net disease in grain amaranth (Amaranthus cruentus L.) in India. Aust Plant Dis 3:129–131

Amaryllis belladonna (Amaryllis, Jersey lily, Naked lily)

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Ramakrishnan G, Ranganathan K, Murugesan S, Sarojini Damodaran AP, Kandaswamy TK (1971) A mosaic disease of Amaranthus gangeticus. Madras Agric J 58:679–683 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Segundo E, Lesemann DE, Martin G, Carmona MP, Cuadrado IM, Velasco L, Janssen D (2007) Amaranthus leaf mottle virus: 30 -end RNA sequence proves classification as distinct virus and reveals affinities within the genus Potyvirus. Eur J Plant Pathol 117:81–87 Shargil D, Smith E, Lachman O, Reingold V, Darzi E, Tam Y, Dombrovsky A (2017) New weed hosts for Cucumber green mottle mosaic virus in wild Mediterranean vegetation. Eur J Plant Pathol 148:473–480 Sharma PN, Chowfla SC (1986) Studies on a mosaic disease of amaranthus (Amaranthus caudatus L.) in Himachal Pradesh. Indian J Mycol Plant Pathol 16(3):349–350 Sharma A, Kulshrestha S (2014) First report of Amaranthus sp. as natural host of Capsicum chlorosis virus in India. Virus Dis 25:412–413 Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct Tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207 Srivastava A, Raj SK, Kumar S, Snehi SK, Kulshreshtha A, Hallan V, Pande SS (2013) Molecular identification of Ageratum enation virus, betasatellite and alphasatellite molecules isolated from yellow vein diseased Amaranthus cruentus in India. Virus Genes 47:584–590 Srivastava A, Kumar S, Raj SK (2014) First report of Ageratum enation virus, betasatellite and alphasatellite causing leaf curl and enation disease of Amaranthus hypochondriacus in India. Plant Dis 98:1285 Srivastava A, Jaidi M, Kumar S, Raj SK, Shukla S (2015) Association of Papaya leaf curl virus with the leaf curl disease of grain amaranth (Amaranthus cruentus L.) in India. Phytoparasitica 43:97–101 Taiwo MA (1988) Studies on a virus disease of Amaranthus hybridus L. in Nigeria. Int J Trop Plant Dis 6:195–200 Vicchi V, Bellardi MG (1988) The role of weeds in the epidemiology of gladiolus viruses and MLO. Acta Hortic 234:371–378 Webster CG, Kousik CS, Roberts PD, Rosskopf EN, Turechek WW, Adkins S (2011) Cucurbit yellow stunting disorder virus detected in pigweed in Florida. Plant Dis 95:360 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Ying ZT, Davis MJ (2000) Partial characterization and host range of Tomato yellow leaf curl virus in South Florida. Proc Florida State Hortic Soc 113:185–190

Amaryllis belladonna (Amaryllis, Jersey lily, Naked lily) Family: Amaryllidaceae

Ornamental

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Amaryllis belladonna was reported from India (Singh et al. 2009). The virus-infected amaryllis plants exhibit foliar symptoms characterized by severe chlorotic stripes, mosaic and yellow mottling, stunting of the spikes, color breaking of the flowers, and reduction of their size. The virus is transmitted by aphid vectors in a non-persistent manner and also by mechanical sap inoculation. For more details of CMV, refer to Cucumis sativus.

Nerine yellow stripe virus Taxonomic position Genus: Potyvirus

(NeYSV)

Family: Potyviridae

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Ambrosia spp. (Ragweed)

NeYSV infection in plants of Amaryllis belladonna was reported from the UK, the Netherlands, Australia, USA, and New Zealand (Pearson et al. 2009; Guaragna et al. 2013). The infected amaryllis plants exhibited mosaic symptoms on the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap inoculation. For more details of NeYSV, refer to Nerine spp.

References Guaragna MA, Lamborn J, Groth-Helms D, Juszczak S, Mollov D, Lockhart B, van ST, Hammond J, Jordan R (2013) First report of Nerine yellow stripe virus in Amaryllis in the United States. Plant Dis 97(10):1389 Pearson MN, Cohen D, Cowell SJ, Jones D, Blouin A, Lebas BSM, Shiller JB, Clover GRG (2009) A survey of viruses of flower bulbs in New Zealand. Australas Plant Pathol 38:305–309 Singh GS, Aminuddin KJ, Srivastava S (2009) First report of Cucumber mosaic virus in Amaryllis belladonna in India. J Plant Pathol 91(S4):107

Ambrosia spp. (Ragweed) Family: Asteraceae

Weed host

Ambrosia asymptomatic virus 1

(ASV1)

Taxonomic position ASV1 is a tentative member of the family Alphaflexiviridae. Geographical distribution ASV1 infection in plants of Ambrosia psilostachya was reported from Oklahoma (USA) (Dutta et al. 2014). Symptoms and host(s) The virus-infected ragweed plants do not exhibit any symptoms. Virion properties and genome The virions are flexible filaments and 12–13 nm in diameter. The virus has a single-stranded RNA genome of about 7408 nt which contains six open reading frames (ORFs) (EU362849–50) (Dutta et al. 2014).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Ambrosia elatior was reported from Hungary (Kazinczi et al. 2001). The virus infected ragweed plants exhibit and mottling symptoms. The virus is transmitted by aphid vectors

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in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

A References Dutta M, Bashir NS, Palmer MW, Melcher U (2014) Genomic characterization of Ambrosia asymptomatic virus 1 and evidence of other Tymovirales members in the Oklahoma tallgrass prairie revealed by sequence analysis. Arch Virol 159:1755–1764 Kazinczi G, Horvath J, Takacs A (2001) Role of weeds in the epidemiology of viruses. In: Proceedings of the 5th Slovenian conference on plant protection, Catez ob Savi, 6–8 Mar 2001, pp 222–226

Ammi majus (Bullwort) Family: Apiaceae

Medicinal plant

Ammi majus latent virus

(AMLV)

Taxonomic position AMLV is a tentative member of the genus Potyvirus and family Potyviridae. Geographical distribution AMLV infection in plants of Ammi majus was reported from Japan (Ueda and Kim, unpublished AB361564). Symptoms and host(s) The virus-infected bullwort plants do not exhibit any symptoms. Transmission The virus is presumably transmitted by aphid vectors. Virion properties and genome The virions are flexuous filaments. The genome is a single molecule of positive-sense single-stranded RNA. A partial polyprotein gene sequence of 1696 nt is available (AB361564) (Wylie et al. 2017).

Apium virus Y

(ApVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

ApVY infection in plants of Ammi majus was reported from Florida (Baker et al. 2008; Adkins et al. 2011). The virus-infected bullwort plants exhibit mosaic, vein-clearing, interveinal chlorosis, rugosity, and deformation symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ApVY, refer to Apium graveolens.

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Amomum subulatum (Large cardamom)

Bidens mottle virus Taxonomic position Genus: Potyvirus

(BiMoV)

Family: Potyviridae

BiMoV infection in plants of Ammi majus was reported from Florida (Baker et al. 2008; Adkins et al. 2011). The virus-infected bullwort plants exhibit mosaic, vein-clearing, interveinal chlorosis, rugosity, and deformation symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BiMoV, refer to Bidens spp.

Clover yellow vein virus Taxonomic position Genus: Potyvirus

(ClYVV)

Family: Potyviridae

ClYVV infection in plants of Ammi majus was reported from Florida (Irey et al. 2006; Adkins et al. 2011). The virus-infected bullwort plants exhibit mosaic, vein-clearing, interveinal chlorosis, and leaf rugosity. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ClYVV, refer to Trifolium spp.

References Adkins S, Baker CA, Jones L, Irey MS, Rosskopf EN (2011) Identification of three potyviruses in Ammi majus in Florida. Acta Hortic 901:133–137 Baker CA, Rosskopf EN, Irey MS, Jones L, Adkins S (2008) Bidens mottle virus and Apium virus Y identified in Ammi majus in Florida. Plant Dis 92:975 Irey M, Adkins S, Baker CA (2006) Clover yellow vein virus identified in Ammi majus in Florida. Plant Dis 90:380 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Amomum subulatum (Large cardamom) Family: Zingiberaceae

Spice crop

Cardamom bushy dwarf virus Taxonomic position Genus: Babuvirus

(CdBDV)

Family: Nanoviridae

Geographical distribution CdBDV infection in plants of Amomum subulatum was reported from India (Varma and Capoor 1964; Mandal et al. 2004; Vijayan et al. 2014; Ghosh et al. 2016a). Earlier this virus disease was popularly known as “Foorkey” disease.

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Symptoms and host(s) The virus-infected large cardamom plants produce many stunted shoots which fail to produce flowers. The leaves are small, slightly curved, and pale green in color. Infection induces excessive sprouting and formation of bushy dwarf clumps develop at the base of mother plants that gradually die. Transmission The virus is transmitted by the aphid vectors Micromyzus kalimpongensis and Pentalonia nigronervosa in a persistent non-propagative manner (Ghosh et al. 2016b). The virus is not mechanically saptransmissible. The use of infected planting material is the primary mode of virus spread (Basu and Ganguly 1968). Virion properties and genome The virions are non-enveloped and spherical of about 17–20 nm in diameter with icosahedral symmetry. The genome is nine positive-sense circular single-stranded (ss) DNAs (NC_038808 to NC_038816): DNA-R, 1102 nt (JX867550); DNA-U3, 1088 nt (KF435143); DNA-S, 1086 nt (JX867540); DNA-M, 1083 nt (JX569847); DNA-C, 1028 nt (JX867548); DNA-N, 1097 nt (JX867546); DNA-U1, 1080 nt (KF435145); DNA-U2, 1079 nt (KF435146); and DNA-U4, 1106 nt (KF435147). Each of the ssDNA segments have a common stem-loop region and is encapsidated in a separate particle (Mandal et al. 2008, 2013; Hu and Yeh 2011).

Large cardamom chirke virus

(LCCV)

Taxonomic position LCCV is a tentative member of the genus Macluravirus and family Potyviridae. Geographical distribution LCCV infection in plants of Amomum subulatum was reported from India (Raychaudhuri and Chatterjee 1958; Raychaudhuri and Ganguly 1965; Mandal et al. 2012; Vijayanandraj et al. 2013; Vijayan et al. 2014). Earlier this virus disease was popularly known as “Chirke” disease. Symptoms and host(s) The virus-infected large cardamom plants exhibit mosaic streaks on the leaves which gradually coalesce and eventually turn brown. The leaves ultimately dry up and wither. Flowering in diseased plants is reduced (Raychaudhuri and Chatterjee 1965). Transmission The virus is transmitted by aphid vectors, Rhopalosiphum maidis, Brachycaudus helichrysi, P. nigronervosa, Sitobion avenae, and Myzus persicae in a non-persistent manner (Mandal et al. 2012; Ghosh et al. 2016b). The virus is sap-transmissible to the popular large cardamom cultivars Golsey, Ramsey, Swaney, and Varlangey. The use of virus-infected planting material is the primary mode of virus spread. Virion properties and genome The virions are flexuous filaments measuring 625–650 nm in length and 12.5 nm in wide. A partial polyprotein gene sequence of 1776 nt is available (JN257715) (Mandal et al. 2012; Wylie et al. 2017).

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Amorphophallus paeoniifolius (Elephant foot yam, Konjac)

References Basu AN, Ganguly B (1968) A note on transmission of ‘Foorkey disease’ of large cardamom by the aphid, Micromyzus kalimpongensis Basu. Indian Phytopathol 21:127 Ghosh A, Das A, Vijayanandraj S, Mandal B (2016a) Cardamom bushy dwarf virus infection in large cardamom alters plant selection preference, life stages, and fecundity of aphid vector, Micromyzus kalimpongensis (Hemiptera: Aphididae). Environ Entomol 45(1):178–184 Ghosh A, Das A, Lepcha R, Mandal B (2016b) Identification, distribution and temporal occurrence of aphids infesting large cardamom and their efficiency in transmitting large cardamom viruses in northeastern sub-Himalayan region. Australas Plant Pathol 45:533–536 Hu M-J, Yeh H-H (2011) Babuvirus. Nanoviridae. In: The Springer index of viruses. Springer, New York, pp 953–958. https://doi.org/10.1007/978-0-387-95919-1_148 Mandal B, Mandal S, Pun KB, Varma A (2004) First report of association of Nanovirus with Foorkey disease of large cardamom in India. Plant Dis 88:428 Mandal B, Mandal S, Tripathi NK, Barman AR, Pun KB, Varma A (2008) Sequence analysis of DNAs encoding putative replicase gene of nanovirus from large cardamom affected by foorkey disease. Indian J Virol 19:S-37 Mandal B, Vijayanandraj S, Shilpi S, Pun KB, Singh V, Pant RP, Jain RK, Varadarasan S, Varma A (2012) Disease distribution and characterisation of a new macluravirus associated with chirke disease of large cardamom. Ann Appl Biol 160:225–236 Mandal B, Shilpi S, Barman AR, Mandal S, Varma A (2013) Nine novel DNA components associated with the foorkey disease of large cardamom: evidence of a distinct babuvirus species in Nanoviridae. Virus Res 178(2):297–305 Raychaudhuri SP, Chatterjee SN (1958) A preliminary note on the occurrence of a new virus disease of large cardamom (Amomum subulatum Roxb.) in Darjeeling District. In: Proceedings of mycological research worker’s conference, ICAR, held at Shimla from the 20th to 23rd June 1958, pp 174–176 Raychaudhuri SP, Chatterjee SN (1965) Transmission of chirke disease of large cardamom by aphid species. Indian J Entomol 27:272–276 Raychaudhuri SP, Ganguly B (1965) Further studies on chirke diseases of large cardamom (Amomum subulatum Roxb.). Indian Phytopathol 18:373–377 Varma PM, Capoor SP (1964) Foorkey disease of large cardamom. Indian J Agrl Sci 34:56–62 Vijayan AK, Gudade BA, Deka TN, Chhetri P (2014) Status of viral diseases of large cardamom (Amomum subulatum Roxb.) and its management in Sikkim and Darjeeling, West Bengal. J Mycol Plant Pathol 44:438–441 Vijayanandraj S, Yogita M, Das A, Ghosh A, Mandal B (2013) Highly efficient immunodiagnosis of Large cardamom chirke virus using the polyclonal antiserum developed against Escherichia coli expressed coat protein. Indian J Virol 24:227–234 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Amorphophallus paeoniifolius (Elephant foot yam, Konjac) Synonyms Amorphophallus konjac Family: Araceae

Dasheen mosaic virus Taxonomic position Genus: Potyvirus

Tuber crop

(DsMV)

Family: Potyviridae

Amorphophallus paeoniifolius (Elephant foot yam, Konjac)

121

DsMV infection in plants of Amorphophallus paeoniifolius was reported from India (Pandit et al. 2001; Ahlawat et al. 2003; Nelson 2008; Babu et al. 2011; Babu and Hegde 2014; Kamala and Makeshkumar 2015). The virus-infected konjac plants exhibit symptoms of chlorotic spotting, leaf mosaic, mottling, puckering, leaf deformation, and shoe stringing symptoms, and infected plants are stunted. The virus is experimentally transmitted by infected sap and, in nature, by several widely distributed aphid species, including Myzus persicae and Aphis gossypii in a non-persistent manner. For more details of DsMV, refer to Colocasia esculenta.

Konjac mosaic virus

(KoMV)

Synonyms Japanese hornwort mosaic virus; Zantedeschia mosaic virus (ZaMV) Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution KoMV infection in plants of Amorphophallus paeoniifolius was first reported from Japan (Shimoyama et al. 1992). The virus spreads in Japan, Taiwan, China, and India (Padmavathi et al. 2011, 2013). Symptoms and host(s) The virus-infected konjac plants exhibit mosaic symptoms. The natural host range of this virus is Amorphophallus paeoniifolius, Colocasia esculenta, Philodendron oxycardium, P. selloum, P. verrucosum, and a few non-aroids that belong to families Chenopodiaceae and Amaranthaceae. Transmission The virus is transmitted by an aphid vector Aphis gossypii in a non-persistent manner. The virus is mechanically sap-transmissible. The virus is transmitted through the use of virus-infected corms. Virion properties and genome The virions are non-enveloped and flexuous filaments 680–900 nm long and 13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9544 nt excluding 30 poly(A) tail (AB219545 = NC_007913). It encodes 350 kDa polyprotein with 3087 amino acid residues (Shimoyama et al. 1992; Nishiguchi et al. 2006; Wylie et al. 2017).

References Ahlawat YS, Pant RP, Bhagawati KN (2003) Association of Carla- and Poty-viruses with mosaic disease of elephant foot yam. Indian Phytopathol 56:300–301 Babu B, Hegde V (2014) Molecular characterization of Dasheen mosaic virus isolates infecting edible aroids in India. Acta Virol 58(1):34–42 Babu B, Hegde V, Makeshkumar T, Jeeva ML (2011) Characterisation of the coat protein gene of Dasheen mosaic virus infecting Elephant foot yam. J Plant Pathol 93(1):199–203 Kamala S, Makeshkumar T (2015) Rapid and sensitive detection of Dasheen mosaic virus infecting elephant foot yam by reverse transcription loop mediated isothermal amplification of coat protein gene. J Virol Methods 222:106–109 Nelson SC (2008) Dasheen mosaic of edible and ornamental aroids. Plant Disease CTAHR. Available at http://www.ctahr. hawaii.edu/oc/freepubs/pdf/PD-44.pdf

A

122

Amsinckia menziesii (Common fiddleneck)

Nishiguchi M, Yamasaki S, Lu XZ, Shimoyama A, Hanada K, Sonada S, Shimon M, Sakai J, Mikoshiba Y, Fujisawa I (2006) Konjac mosaic virus: the completed nucleotide sequences of the genomic RNA and its comparison with other potyviruses. Arch Virol 151:1643–1650 Padmavathi M, Srinivas KP, Subba Reddy CV, Ramesh B, Navodayam K, Krishnaprasadji J, Babu Ratan P, Sreenivasulu P (2011) Konjac mosaic virus naturally infecting three aroid plant species in Andhra Pradesh, India. J Phytopathol 159:133–135 Padmavathi M, Srinivas KP, Hema M, Sreenivasulu P (2013) First report of Konjac mosaic virus in elephant foot yam (Amorphophallus paeoniifolius) from India. Australas Plant Dis Notes 8:27–29 Pandit MK, Mukhopadhyay S, Devonshire BJ, Jones P (2001) First report of Dasheen mosaic virus in Elephant foot yam in India. Plant Pathol 50:802 Shimoyama J, Kameya-Iwaki M, Hanada K, Gunji T (1992) Konjak mosaic virus, a new potyvirus infecting konjac, Amorphophallus konjac. Ann Phytopath Soc Japan 58:706–712 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Amsinckia menziesii (Common fiddleneck) Family: Boraginaceae

Beet curly top virus Taxonomic position Genus: Curtovirus

Weed host

(BCTV)

Family: Geminiviridae

BCTV infection in plants of Amsinckia menziesii was reported from California (USA) (Creamer et al. 1996). The virus is transmitted by a leafhopper vector, Circulifer tenellus, in a persistent (circulative, non-propagative) manner. The virus is not transmitted by mechanical inoculation. For more details of BCTV, refer to Beta vulgaris.

References Creamer R, Lugue-Williams M, Howo M (1996) Epidemiology and incidence of Beet curly top geminivirus in naturally infected weed hosts. Plant Dis 80:533–535

Anagyris foetida (Purging trefoil) Family: Fabaceae

Medicinal plant

Anagyris vein yellowing virus Taxonomic position Genus: Tymovirus

(AVYV)

Family: Tymoviridae

Ananas comosus (Pineapple)

123

Geographical distribution AVYV infection in plants of Anagyris foetida was reported from Southern Italy (Rana et al. 1988).

A Symptoms and host(s) The virus-infected purging trefoil plants exhibit vein yellowing symptoms. Transmission The virus is transmitted by beetle vectors in a semi-persistent manner. The virus has been mechanically transmitted to several plant species belonging to the Leguminosae and the Solanaceae. Virion properties and genome The virions are non-enveloped, isometric capsid with T = 3 icosahedral symmetry, about 30 nm in diameter. The genome consists of a single molecule of positive-sense, ssRNA of 6151 nt (AY751780 = NC_011559). The 30 -terminus has a tRNA-like structure (Koenig et al. 2005; Martelli et al. 2002).

Scrophularia mottle virus Taxonomic position Genus: Tymovirus

(SrMV)

Family: Tymoviridae

SrMV infection in plants of Anagyris foetida was reported by Rana et al. (1988). The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of SrMV, refer to Scrophularia spp.

References Koenig R, Pleij CWA, Lesemann DE, Loss S, Vetten HJ (2005) Molecular characterization of isolates of Anagyris vein yellowing virus, Plantago mottle virus and Scrophularia mottle virus comparison of various approaches for Tymovirus classification. Arch Virol 150:2325–2338 Martelli GP, Sabanadzovic S, Abou-Ghanem Sabanadzovic N, Edwards MC, Dreher T (2002) The family Tymoviridae. Arch Virol 147(9):1837–1846 Rana GL, Castellano MA, Koenig R (1988) Characterization of a Tymovirus isolated from Anagyris foetida as a strain of Scrophularia mottle virus. J Phytopathol 121:239–249

Ananas comosus (Pineapple) Family: Bromeliaceae

Edible fruit crop

Endogenous pineapple pararetrovirus 1

(EPPV1)

Taxonomic position EPPV1 is a tentative member of the genus Badnavirus and family Caulimoviridae.

124

Ananas comosus (Pineapple)

Geographical distribution EPPV1 infection in plants of Ananas comosus was reported from Cuba, Australia, and Hawaii (Hernandez-Rodriguez et al. 2013). Symptoms and host(s) The virus has been detected in pineapple plants showing mealybug wilt disease symptoms (foliar reddening, leaves with tips curved down, and dieback). However, the specific symptom of the virus have not been confirmed through a transmission test. Virion properties and genome There is no evidence of encapsidation of genome in virions. The genome consists of a circular dsDNA. Partial polyprotein gene sequences are available (EU377674, GQ395780, JQ390622) (Gambley et al. 2008; Hernandez-Rodriguez et al. 2013).

Pineapple bacilliform CO virus Taxonomic position Genus: Badnavirus

(PBCOV)

Family: Caulimoviridae

Geographical distribution PBCOV infection in plants of Ananas comosus was reported from Hawaii, China, Australia, and Cuba (Gambley et al. 2008; Wu et al. 2010; Sether et al. 2012; Hernandez-Rodriguez et al. 2013). Symptoms and host(s) The virus-infected pineapple plants exhibit symptoms of mosaic, chlorosis, veinbanding, and stunting. Transmission The virus is transmitted by gray pineapple mealybugs, Dysmicoccus neobrevipes and Planococcus citri, in a semi-persistent manner (Sether et al. 2012). Virion properties and genome The virions are bacilliform in shape with a diameter of 30 nm, and its modal particle length is 130 nm (Olszewski and Lockhart 2011; Bhat et al. 2016). The genome is a circular dsDNA. Complete genome sequence are available from Hawaii, 7451 bp (GQ398110) (Sether et al. 2012), and China, 7543 bp (GUi21676 = NC_014648) (Wu et al. 2010) (Olszewski and Lockhart 2011; Bhat et al. 2016).

Pineapple bacilliform ER virus Taxonomic position Genus: Badnavirus

(PBERV)

Family: Caulimoviridae

Geographical distribution PBERV infection in plants of Ananas comosus var. erectifolius was reported from Australia (Gambley et al. 2008).

Ananas comosus (Pineapple)

125

Symptoms and host(s) No definite symptom has been associated with infection by this virus. Transmission The virus is transmitted by the gray pineapple mealybug, Dysmicoccus neobrevipes, in a semipersistent manner (Gambley et al. 2008). Virion properties and genome The virions are bacilliform in shape with a diameter of 30 nm, and its modal particle length is 130 nm. The complete genome sequence of PBERV is not available. The partial genome sequence that containing the RT/RNase H region 1510 bp (EU377672, EU377673) showed a phylogenetic affinity with badnaviruses (Gambley et al. 2008; Olszewski and Lockhart 2011; Bhat et al. 2016).

Pineapple chlorotic leaf streak virus

(PCLSV)

Taxonomic position PCLSV is a tentative member of the family Rhabdoviridae. Geographical distribution PCLSV infection in plants of Ananas comosus was reported from Brazil (Kitajima et al. 1975). Symptoms and host(s) The virus-infected pineapple plants exhibit conspicuous chlorotic leaf streak symptoms. Transmission The virus is not mechanically sap-transmissible. Virion properties and genome The virions are bacilliform or rhabdo- or bullet-shaped with a clear modal length of 200–250 nm and 60–70 nm wide (Kitajima et al. 1975).

Pineapple mealybug wilt associated virus

(PMWaV)

Synonyms Pineapple wilt-associated virus Taxonomic position PMWaV is a tentative member of the genus Ampelovirus and family Closteroviridae. Geographical distribution PMWaV infection was first reported in plants of Ananas comosus cv. Smooth Cayenne from Hawaii by Carter (1933). The virus spreads in Indonesia, Malaysia, Sabah, the Philippines, Taiwan, Mauritius, South Africa, Sri Lanka, Australia, India, Jamaica, Fiji, Cuba, Central and South America, and the USA (Hawaii) (Westgate 1945; Carter 1973; Singh and Sastry 1974; German et al. 1992; Wakman et al. 1995; Hu et al. 1996).

A

126

Ananas comosus (Pineapple)

Symptoms and host(s) On pineapple the virus symptoms are characterized by initial leaf-tip dieback and reddening along the leaf length. This is followed by a progressive change of leaf color from red to pink, inward reflexing of the leaf margin, loss of rigidity, and the eventual collapse of the leaves along most of their length. Necrosis of leaf tips, wilting, and eventual plant death occur (Gunasinghe 1989; Rohrbach et al. 1988). Earlier reports of pineapple mealybug wilt-associated virus may have represented various ampeloviruses, which based on nucleotide sequence, genome organization, and phylogenetic analysis of pineapple mealybug wilt-associated viruses are now differentiated as PMWaV-1, PMWaV-2, PMWaV-3, and PMWaV-4. Transmission The virus is transmitted by mealybug vectors Dysmicoccus brevipes, D. neobrevipes, and Pseudococcus longispinus in a semi-persistent manner. Even a single specimen of the mealybug can transmit the virus, but for maximum virus transmission, a minimum of 20 mealybugs per plant are required (Sether et al. 1998). Sether et al. (1998) reported that all stages of D. neobrevipes acquired the virus. The virus is not transmitted by mechanical inoculation (Carter 1934; Singh and Sastry 1974). Virion properties and genome The virions are flexuous filaments with a clear modal length of 1200–1500 nm and width of 12 nm (Gunasinghe and German 1986). The genome is a single-stranded RNA (Ullman et al. 1989; Borroto et al. 1998).

Pineapple mealybug wilt-associated virus 1 Taxonomic position Genus: Ampelovirus

(PMWaV-1)

Family: Closteroviridae

Geographical distribution PMWaV-1 infection in plants of Ananas comosus was reported from Australia, China, Taiwan, and Hawaii (Hu et al. 1997; Sether and Hu 2001; Liao et al. 2008; Gambley et al. 2009; Shen et al. 2009; Gambley et al. 2010; Hernandez-Rodriguez et al. 2014; Alvarez et al. 2015; Yu et al. 2015). Symptoms and host(s) The virus causes latent infection in pineapple. In Hawaii, a high incidence of PMWaV-1, up to 100%, was detected in asymptomatic pineapple. The study on the effect of PMWaV-1 infection on fruit yield of pineapple revealed that PMWaV-1 infection had no significant effects on fruit weight in the main crop, whereas PMWaV-1 infection significantly reduced the fruit size in the ratoon crop. PMWaV-1 infection and reduced irrigation had a combined effect resulting in a 13.4% yield loss in the ratoon crop (Sether and Hu 2001). Transmission The virus is transmitted semi-persistently by mealybugs (Dysmicoccus spp.) (Sether et al. 1998). The virus is not transmitted by mechanical sap-inoculation, and not by seed. Virion properties and genome The virions are flexuous filaments measuring 1250–1300  12–15 nm in dimension (Liao et al. 2008). The genome is a monopartite, linear, single-stranded, positive-sense RNA of 13,071 nt (AF414119 =

Ananas comosus (Pineapple)

127

NC_010178) with 7 open-reading frames (ORFs) (Maliogka et al. 2011; Melzer et al. 2008; Dey et al. 2015).

A Pineapple mealybug wilt-associated virus 2 Taxonomic position Genus: Ampelovirus

(PMWaV-2)

Family: Closteroviridae

Geographical distribution PMWaV-2 was first detected in a diseased Ananas comosus plants from Ciego de Avila (Cuba) in 1998. The virus spreads in Taiwan, Australia, Thailand, Hawaii, and Cuba (Melzer et al. 2001; Sether and Hu 2002; Borroto-Fernandez et al. 2007; Gambley et al. 2009, 2010; Shen et al. 2009). Symptoms and host(s) The virus-infected pineapple plants show typical symptoms of foliar reddening, leaves with tips curved down, and dieback. Transmission The virus is transmitted by the gray pineapple mealybug (GPM) vector Dysmicoccus neobrevipes in a semi-persistent manner. Some GPM were able to acquire the virus within 36 h. Transmission efficiency of GPM in groups of 25 was 100% following a 72-h acquisition access period (AAP). GPM remained viruliferous for up to 3 days after a 3-day AAP when serially transferred to virus-free pineapple plants at 24-h intervals over a 7-day period (Subere et al. 2011). The virus is not transmitted by mechanical sapinoculation, and not by through seed. Virion properties and genome The virions are filamentous, non-enveloped, and very flexuous, measuring 1400–2000 nm long and 10–13 nm in diameter. The genome is a monopartite, linear, single-stranded, positive-sense RNA of 14,861 nt (AF283103) (Melzer et al. 2001; Borroto-Fernandez et al. 2007; Maliogka et al. 2011; Dey et al. 2015).

Pineapple mealybug wilt-associated virus 3 Taxonomic position Genus: Ampelovirus

(PMWaV-3)

Family: Closteroviridae

Geographical distribution PMWaV-3 infection in plants of Ananas comosus was reported from Thailand, Australia, Hawaii, and Cuba (Gambley et al. 2009, 2010; Shen et al. 2009; Hernandez et al. 2010). Symptoms and host(s) The virus-infected pineapple plants show typical symptoms of foliar reddening, leaves with tips curved down, and dieback.

128

Ananas comosus (Pineapple)

Transmission The virus is transmitted by mealybug vectors Dysmicoccus brevipes and D. neobrevipes in a semipersistent manner (Sether et al. 2005). The virus is not transmitted by mechanical sap-inoculation, and not through seed. Virion properties and genome The virions are filamentous, non-enveloped, and very flexuous, measuring 1400–2000 nm long and 10–13 nm in diameter. The genome is a monopartite, linear, single-stranded, positive-sense RNA of 13,500 nt (DQ399259) (Sether et al. 2009; Maliogka et al. 2011).

Tomato spotted wilt orthotospovirus

(TSWV)

Synonyms Pineapple yellow spot virus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

TSWV was originally reported as Pineapple yellow spot virus infecting Ananas comosus in Hawaii (Illingworth 1931; Linford 1932). The virus is probably distributed worldwide wherever Ananas comosus is grown. The disease takes its name from the appearance of the early symptoms on leaves, namely, a small, round, yellowish spot “eye” on the upper surface of the pineapple leaves; these later enlarge downward toward the leaf base, increasing into irregular streaks. One of the symptoms on the fruit is a blackened, dry cavity in the side of the fruit due to one or more “eyes” having died. This is known as “dead eye.” Infection of very young fruit results in an irregular arrangement of fruitlets as some fail to develop. Attacked fruits may also fail to develop a crown. Thrips feeding on the crown of fruits result in concentric ring patterns developing on crown leaves. These spots enlarge, and the infection spreads into the fruit itself, by which time the crown will often have dried out. The flesh of fruits thus infected will be discolored and necrotic below the butt of the crown. The whole fruit may eventually become affected. The virus has a very wide host range. The virus is transmitted by thrips vectors Thrips tabaci, T. setosus, T. parmi, Frankliniella schultzei, F. occidentalis, F. fusca, and Scirtothrips dorsalis in a persistent-propagative manner (Linford 1932). The virus is also mechanically sap-transmissible. The virus is not pollen and seed-transmitted. For more details of TSWV, refer to Solanum lycopersicum.

References Alvarez RA, Martin RR, Quito-Avila DF (2015) First report of Pineapple mealybug wilt associated virus-1 in Ecuador. New Dis Rep 31:15 Bhat AI, Hohn T, Selvarajan R (2016) Badnaviruses: the current global scenario. Viruses 8(6):177 Borroto EG, Cintra M, Gonsalvez J, Borroto C (1998) First report of closterovirus-like particles associated with pineapple plants (Ananas comosus cv. Smooth Cayenne) affected with pineapple mealybug wilt in Cuba. Plant Dis 82:263 Borroto-Fernandez EG, Torres-Acosta JA, Laimer M (2007) RT-PCR detection and protein-protein interaction of viral components of Pineapple mealybug wilt-associated virus 2 in Cuba. J Plant Pathol 89:435–439 Carter W (1933) The pineapple mealybug, Pseudococcus brevipes (Ckl.) and wilt of pineapples. Phytopathology 23:207–242 Carter W (1934) The mealybug wilt and green spots in Jamaica and Central America. Phytopathology 24:424–426

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Carter W (1973) Mealybug wilt of pineapple. FAO Plant Protect Bull 21(5):115 Dey KK, Borth WB, Melzer MJ, Wang M-L, Hu JS (2015) Analysis of Pineapple mealybug wilt associated virus -1 and -2 for potential RNA silencing suppressors and pathogenicity factors. Viruses 7:969–995 Gambley CF, Geering ADW, Steele V, Thomas JE (2008) Identification of viral and non-viral reverse transcribing elements in pineapple (Ananas comosus), including members of two new badnavirus species. Arch Virol 153:1599–1604 Gambley CF, Geering ADW, Thomas JE (2009) Development of an immunomagnetic capture-reverse transcriptase-PCR assay for three pineapple ampeloviruses. J Virol Methods 155:187–192 Gambley CF, Steele V, Geering ADW, Thomas JE (2010) The diversity of ampeloviruses and badnaviruses in Australian pineapples and their association with mealybug wilt of pineapple (Ananas comosus). In: 11th international plant virus epidemiology symposium held at Cornell University, New York, Abstract German TL, Ullman DE, Gunashinghe UB (1992) Mealybug wilt of pineapple. Adv Dis Vector Res 9:241–259 Gunasinghe UB (1989) Characterization of new virus isolated from pineapple. PhD thesis, University of Hawaii, Honolulu Gunasinghe UB, German TL (1986) Association of virus particles with mealybug-wilt of pineapple (Abstr.). Phytopathology 76:1073 Hernandez L, Ramos PL, Rodriguez M, Pena I, Perez JM (2010) First report of Pineapple mealybug wilt associated virus3 infecting pineapple in Cuba. New Dis Rep 22:18 Hernandez-Rodriguez L, Ramos-Gonzalez PL, Garcia-Garcia G, Javer Higginson E, Zamora-Rodriguez V (2013) First report of Pineapple bacilliform comosus virus (PBCoV) and endogenous Pineapple pararetrovirus-1 (ePPRV-1) in pineapple plants in Cuba. New Dis Rep 28:2 Hernandez-Rodriguez L, Ramos-Gonzalez PL, Garcia-Garcia G, Zamora V, Peralta-Martin AM, Peña I, Perez JM, Ferriol X (2014) Geographic distribution of mealybug wilt disease of pineapple and genetic diversity of viruses infecting pineapple in Cuba. Crop Prot 65:43–50 Hu JS, Sether DM, Ullman DE (1996) Detection of pineapple Closterovirus in pineapple plants and mealybugs using monoclonal antibodies. Plant Pathol 45(5):829–836 Hu JS, Sether DM, Liu XP, Wang M, Zee F, Ullman D (1997) Use of a tissue blotting immunoassay to examine the distribution of pineapple closterovirus in Hawaii. Plant Dis 81:1150–1154 Illingworth JF (1931) Yellow spot of pineapples in Hawaii. Phytopathology 21:865–880 Kitajima EW, Giacomelli EJ, Costa AS, Costa CL, Cupertino FP (1975) Bacilliform particles associated with chlorotic leaf streak of Giant pineapple (Ananas comosus L.). Merrill Phytopath Z 82:83–86 Liao JY, Hu CC, Chang CA, Deng TC (2008) The preliminary identification of Pineapple mealybug wilt-associated virus1 on pineapple in Taiwan. J Taiwan Agric Res 57:1–14 Linford B (1932) Transmission of the Pineapple yellow-spot virus by Thrips tabaci. Phytopathology 22: 301–324 Maliogka V, Dovas C, Katis N (2011) Ampelovirus. Closteroviridae. In: The Springer index of viruses. Springer, New York, pp 317–326. https://doi.org/10.1007/978-0-387-95919-1_49 Melzer MJ, Karasev AV, Sether DM, Hu JS (2001) Nucleotide sequence, genome organization, and phylogenetic analysis of Pineapple mealybug wilt associated virus 2. J Gen Virol 82:1–7 Melzer MJ, Sether DM, Karasev AV, Hu JS (2008) Complete nucleotide sequence and genome organization of Pineapple mealybug wilt-associated virus-1. Arch Virol 153:707–714 Olszewski NE, Lockhart B (2011) Badnavirus. Caulimoviridae. In: The Springer index of viruses. Springer, New York, pp 263–269. https://doi.org/10.1007/978-0-387-95919-1_40 Rohrbach KG, Beardsley JW, German TL, Reimer N, Sanford WG (1988) Mealybug wilt, mealybugs and ants on pineapple. Plant Dis 72:558–565 Sether DM, Hu JS (2001) The impact of Pineapple mealybug wilt-associated virus-1 and reduced irrigation on pineapple yield. Australas Plant Pathol 30:31–36 Sether DM, Hu JS (2002) Yield impact and spread of Pineapple mealybug wilt associated virus-2 and mealybug wilt of pineapple in Hawaii. Plant Dis 86:867–874 Sether D, Ullman D, Hu J (1998) Transmission of Pineapple mealybug wilt-associated virus by two species of mealybug (Dysmicoccus spp.). Phytopathology 88:1224–1230 Sether DM, Melzer MJ, Busto J, Zee F, Hu JS (2005) Diversity and mealybug transmissibility of ampeloviruses in pineapple. Plant Dis 89:450–456 Sether DM, Melzer MJ, Borth WB, Hu JS (2009) Genome organization and phylogenetic relationship of Pineapple mealybug wilt-associated virus-3 with family Closteroviridae members. Virus Genes 38:414–420 Sether DM, Melzer MJ, Borth WB, Hu JS (2012) Pineapple bacilliform CO virus: diversity, detection, distribution, and transmission. Plant Dis 96:1798–1804 Shen BN, Zheng YX, Chen WH, Chang TY, Ku H-M, Jan F-J (2009) Occurrence and molecular characterization of three pineapple mealybug wilt-associated viruses in pineapple in Taiwan. Plant Dis 93:196 Singh SJ, Sastry KSM (1974) Wilt of pineapple - a new virus disease in India. Indian Phytopath 27:298–303

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130

Andrographis paniculata (Green chirayta, Kalmegh)

Subere CVQ, Sether DM, Borth WB, Melzer MJ, Hu JS (2011) Transmission characteristics of Pineapple mealybug wilt associated virus-2 by the grey pineapple mealybugs Dysmicoccus neobrevipes in Hawaii. Acta Hortic (ISHS) 902:393–399 Ullman DE, German TL, Gunasinghe UB, Ebusu RH (1989) Etiology of Closterovirus-like particles associated with mealybug wilt of pineapple. Phytopathology 79:1341–1345 Wakman W, Teakle DS, Thomas JE, Dietzgen RG (1995) Presence of a clostero like virus and bacilliform virus in pineapple in Australia. Aust J Agric Res 46:947–958 Westgate PJ (1945) Mealybug wilt of pineapple in South Florida. Proc Fla State Hortic Soc 58:194–196 Wu L, Ruan X, Shen W, Tan Y, Zhai G, Li H (2010) Sequencing and analysis of the complete genomic sequence of pineapple bacilliform comosus virus. Sci Agric Sin 43:1969–1976 Yu N, Luo Z, Fan H, Zhang Z, Li X, Wang J, Liu Z, He F (2015) Complete genomic sequence of a Pineapple mealybug wilt-associated virus-1 from Hainan Island, China. Eur J Plant Pathol 141:611–615

Andrographis paniculata (Green chirayta, Kalmegh) Family: Acanthaceae

Medicinal plant

Andrographis yellow vein leaf curl virus Taxonomic position Genus: Begomovirus

(AnYVLCV)

Family: Geminiviridae

Geographical distribution AnYVLCV infection in plants of Andrographis paniculata was reported from India (Khan et al. 2015). Symptoms and host(s) The virus-infected kalmegh plants exhibit prominent lethal leaf yellowing symptoms. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm, with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2754 nt (KM359406 = NC_028262) (Briddon 2001; Brown et al. 2015; Khan et al. 2015; Zerbini et al. 2017).

Catharanthus yellow mosaic virus Taxonomic position Genus: Begomovirus

(CatYMV)

Family: Geminiviridae

CatYMV infection in plants of Andrographis paniculata was reported from India (Khan et al. 2015). The virus-infected kalmegh plants initially showed some chlorotic streaks, which later turned completely yellow, ultimately leading to premature death. The virus is transmitted by the whitefly

Andrographis paniculata (Green chirayta, Kalmegh)

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vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of CatYMV, refer to Catharanthus spp.

A Eclipta yellow vein virus Taxonomic position Genus: Begomovirus

(EYVV)

Family: Geminiviridae

EYVV infection in plants of Andrographis paniculata was reported from India (Khan and Samad 2014; Khan et al. 2016). The virus-infected kalmegh plants exhibit symptoms of yellow veins on younger leaves and later upward leaf curling, vein clearing, chlorosis, reduced leaf size, poor inflorescence development, and stunted growth leading to drastic reduction in herb yield. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner. For more details of EYVV, refer to Eclipta prostrata (Syn.) Eclipta alba.

Hollyhock leaf curl virus

(HoLCV)

Synonyms Eclipta yellow vein virus (EYVV) Taxonomic position Genus: Begomovirus

Family: Geminiviridae

EYVV infection in plants of Andrographis paniculata was reported from India (Khan and Samad 2014). The virus-infected kalmegh plants show yellow veins on younger leaves and later upward leaf curling, vein-clearing, chlorosis, reduced leaf size, poor inflorescence, and stunted growth. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Andrographis yellow vein leaf curl betasatellite DNA molecule is associated with EYVV, and consists of 1379 nt (KC967282) (Zhou 2013; Khan and Samad 2014). For more details of HoLCV, refer to Alcea rosea (syn. Althaea rosea).

References Briddon RW (2001) Begomovirus. Geminiviridae. The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Khan A, Samad A (2014) A new isolate of Eclipta yellow vein virus along with a betasatellite associated with yellow vein leaf curl disease of Andrographis paniculata in India. Plant Dis 98:698 Khan A, Saeed ST, Samad A (2015) New record of Catharanthus yellow mosaic virus and a betasatellite associated with lethal leaf yellowing of kalmegh (Andrographis paniculata) in northern India. Plant Dis 99:292 Khan A, Luqman S, Masood N, Singh DK, Saeed ST, Samad A (2016) Eclipta yellow vein virus enhances chlorophyll destruction, singlet oxygen production and alters endogenous redox status in Andrographis paniculata. Plant Physiol Biochem 104:165–173 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zhou X (2013) Advances in understanding begomovirus satellites. Annu Rev Phytopathol 51:357–381

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Anemone coronaria (Anemone)

Anemone coronaria (Anemone) Family: Ranunculaceae

Ornamental

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV-infected Anemone spp. have been reported from the UK, the USA, the Netherlands, Israel, and Greece (Loebenstein et al. 1995; Chen 2003; Tzanetakis 2009; Xanthis et al. 2015). The virus-infected anemone plants exhibit leaves which are conspicuously chlorotic and “broken flowers” with fine white or pale lines and streaks in red and blue petals (Alper et al. 1989). The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Anemone spp. was reported from Italy (Vaira et al. 1993; Gera et al. 1999). The virus-infected anemone plants exhibit foliar ring-spots and foliar necrosis symptoms. The virus is transmitted by a thrips vector, Frankliniella occidentalis, in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of INSV, refer to Impatiens spp.

Ranunculus mild mosaic virus Taxonomic position Genus: Potyvirus

(RanMMV)

Family: Potyviridae

RanMMV was detected in plants of Anemone spp. in the Netherlands (Pham et al. 2011). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of RanMMV, refer to Ranunculus asiaticus.

Ranunculus white mottle ophiovirus Taxonomic position Genus: Ophiovirus

(RWMV)

Family: Aspiviridae

RWMV infection in plants of Anemone coronaria was reported from Italy (Vaira et al. 2000, 2002). The virus-infected anemone plants were observed with symptoms on young leaves showing curling, deformation, necrotic spotting, and stunting. No vector is known for this virus. The virus is mechanically sap-transmissible. For more details of RWMV, refer to Ranunculus asiaticus.

Anemone coronaria (Anemone)

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Raspberry ringspot virus Taxonomic position Genus: Nepovirus

(RpRSV)

A

Family: Secoviridae

RpRSV infection in plants of Anemone coronaria was reported from Italy, Holland, and Germany (Kobayashi 1981; Lisa et al. 2002). Anemone (cv. Whirlwind) plants infected with RpRSV show ringspot and mottling symptoms on the leaves and stunting. The virus is transmitted by the nematode vector Longidorus spp., and there is also a report of RpRSV transmission through members of the genera Paratrichodorus and Xiphinema in a non-persistent manner. The virus is mechanically saptransmissible and also by grafting. The use of vegetative material from virus-infected plants for planting is the major mode of virus spread. For more details of RpRSV, refer to Rubus spp.

Strawberry latent ringspot virus Taxonomic position Genus: Unassigned

(SLRSV)

Family: Secoviridae

SLRSV infection in plants of Anemone coronaria was reported from New Zealand (Tang et al. 2013). The virus-infected anemone plants exhibit symptoms of chlorotic spots on the leaves. The virus is transmitted by nematode vectors (Xiphinema spp.) and also by mechanical sap-inoculation. For more details of SLRSV, refer to Fragaria spp.

Tobacco necrosis virus

(TNV)

Taxonomic position TNV is a tentative member of the genus Necrovirus and family Tombusviridae TNV infection in plants of Anemone coronaria was reported by Alper et al. (1989). The virus apparently occurs symptomlessly in anemone plants and is often detected in roots but more rarely in leaves. The virus is transmitted by the zoospores of the fungus Olpidium brassicae and also by mechanical sapinoculation. For more details of TNV, refer to Nicotiana tabacum.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in plants of Anemone coronaria was reported from the USA, Germany, and the UK (Lockhart and Westendrop 1998; Lisa et al. 2002; Harju et al. 2011). The virus-infected anemone plants exhibit symptoms consisting of chlorotic blotches, chlorotic line patterns, and distortion. The virus is transmitted by nematode vectors and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

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Anemone coronaria (Anemone)

Tobacco ringspot virus

(TRSV)

Synonyms Anemone necrosis virus Taxonomic position Genus: Nepovirus

Family: Secoviridae

TRSV infection in plants of Anemone coronaria was reported from the UK, the Netherlands, Israel, and Italy (Loebenstein et al. 1995). The young leaves and then the petioles of TRSV-infected anemone plants become necrotic, and such plants fail to flower and often die prematurely; however, partially recovered plants had less severe symptoms which included slight leaf mottling, necrosis, and dwarfing. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sapinoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Anemone coronaria was reported from the USA and Italy (Hurtt et al. 1985; Vaira et al. 1993; Daughtrey 1996; Gera et al. 1998). The virus-infected anemone plants exhibit vein-clearing, chlorotic/necrotic spots and rings on the leaves, necrosis of petioles and stems, and severe stunting of plants (Loebenstein et al. 1995). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Turnip mosaic virus

(TuMV)

Synonyms Anemone mosaic virus (AnMV) Taxonomic position Genus: Potyvirus

Family: Potyviridae

The “Anemone mosaic virus” reported by Hollings (1957) was later recognized as a strain of TuMV. The virus-infected Anemone coronaria plants were reported from the Netherlands (Pham et al. 2011). The virus-infected anemone plants were stunted and had slightly mottled and distorted leaves and “broken flowers”; the leaf chlorosis in some plants, however, was very conspicuous. In winter the petioles of infected plants developed necrotic pitting and streaking which eventually resulted in dieback and death of some plants (Alper et al. 1989). The virus is transmitted by aphid vectors in a nonpersistent manner, and also by mechanical sap-inoculation. For more details of TuMV, refer to Brassica rapa.

Anethum graveolens (Dill)

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References Alper M, Levy S, Loebenstein G (1989) Virus diseases in anemones. Hasadeh 65:105 Chen YK (2003) Occurence of Cucumber mosaic virus in ornamental plants and perspectives of transgenic control. PhD thesis, Wageningen University, The Netherlands. p 144 Daughtrey ML (1996) Detection and identification of tospoviruses in greenhouses. Acta Hortic 431:90–98 Gera KA, Cohen KH, Beckelman H, Anav D, Hadr E, Levy E, Koperman A (1998) Identification of Tomato spotted wilt tospovirus in Anemone. Dapey Medaa 12:69–72 Gera A, Kritzman A, Cohen J, Raccah B (1999) First report of Impatiens necrotic spot tospovirus (INSV) in Israel. Plant Dis 83:587 Harju VA, Skelton AL, Monger WA, Forde SMD, Daly M, Nixon T, Lawson R, Bennett S, Mumford RA (2011) A report of viruses, viroids and phytoplasmas found in ornamental plants from 1999–2007: findings from central science laboratory, UK. Acta Hortic 901:223–229 Hollings M (1957) Anemone mosaic – a virus disease. Ann Appl Biol 45:44–61 Hurtt SS, Trees SC, Watterson JC (1985) Tomato spotted wilt virus associated with a lethal disease of Anemone coronaria. Phytopathology 75:1359 Kobayashi T (1981) Raspberry ringspot virus detected in anemones imported from Holland. Res Bull Plant Protect Serv Jpn 5:42 Lisa V, Vaira AM, Milne RG, Lesemann D-E (2002) Virus diseases of Japanese anemone. Acta Hortic 568:185–191 Lockhart BE, Westendrop JA (1998) Anemone - an additional perennial ornamental host of Tobacco rattle virus in the U.S. Plant Dis 82:712 Loebenstein G, Lawson RH, Brunt AA (1995) Virus and virus-like diseases of bulb and flowering crops. Wiley, United Kingdom, p 543 Pham KTK, de Kock MJD, Lemmers MEC, Derks AFLM (2011) Molecular identification of potyviruses infecting bulbous ornamentals by the analysis of coat protein (CP) sequences. Acta Hortic 901:167–172 Tang J, Ward LI, Clover GRG (2013) The diversity of Strawberry latent ringspot virus in New Zealand. Plant Dis 97:662–667 Tzanetakis IE (2009) First report of Cucumber mosaic virus in Anemone sp. in the United States. Plant Dis 93:431 Vaira AM, Roggero P, Luisoni E, Masenga V, Milne RG, Lisa V (1993) Characterization of two tospoviruses in Italy: Tomato spotted wilt and impatiens necrotic spot. Plant Pathol 42:530–542 Vaira AM, Vecchiati M, Lisa V, Milne RG (2000) First report of anemone plants infected by Ranunculus white mottle virus. Plant Dis 84:1046 Vaira AM, Accotto GP, Lisa V, Vecchiati M, Masenga V, Milne RG (2002) Molecular diagnosis of Ranunculus white mottle virus in two ornamental species. Acta Hortic 568:29–33 Xanthis CK, Maliogka VI, Katis NI (2015) First report of Cucumber mosaic virus in Anemone sp. in Greece. J Plant Pathol 97:541

Anethum graveolens (Dill) Family: Apiaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Medicinal plant

(AMV)

Family: Bromoviridae

AMV infection in plants of Anethum graveolens was reported from Hungary and Yemen (Wolf 1972; Alhubaishi et al. 1987). The virus-infected dill plants exhibit mottling, leaf necrosis, dwarfing, and malformation symptoms. The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

A

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Anethum graveolens (Dill)

Carrot red leaf virus

(CtRLV)

Taxonomic position Genus: Polerovirus

Family: Luteoviridae

CtRLV infection in plants of Anethum graveolens was reported from Australian Capital Territory (Waterhouse 1985). The virus-infected dill plants show symptoms of stunting and reddening or yellowing of the leaves. The virus is not mechanically sap-transmissible but is transmitted by the aphid vectors in a circulative, non-propagative manner. For more details of CtRLV, refer to Daucus carota.

Celery mosaic virus

(CeMV)

Synonyms Western celery mosaic virus

Taxonomic position Genus: Potyvirus

Family: Potyviridae

CeMV infection in plants of Anethum graveolens was reported from Northern Italy and Hungary (Wolf 1972; Bellardi and Rubies-Autonell 1998). The virus-infected dill plants exhibit symptoms of narrowing of the leaf lamina and stunting that evolved to yellow or bronze color. The plants appeared bushy and had low seed production. The virus is transmitted by a number of aphid species in a nonpersistent manner, and also by mechanical sap-inoculation to a limited host range. The virus is not transmissible by contact between plants. For more details of CeMV, refer to Apium graveolens.

Dill clover cryptic virus 1

(DCV1)

Taxonomic position DCV1 is a tentative member of the genus Alphapartitivirus and family Partitiviridae

Geographic distribution DCV1 infection in plants of Anethum graveolens var. hortorum was reported from Germany (Lesker and Maiss 2013; Lesker et al. 2013).

Transmission The virus is transmitted through pollen and seed.

Virion properties and genome The virions are isometric, 25–50 nm in diameter. The genome consists of double-stranded RNA of two parts. RNA1 consists of 2013 bp (KF484726) and RNA2 is 1837 bp (KF484727) (Vainio et al. 2018).

Angelica spp.

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Dill cryptic virus 2

(DCV2)

Taxonomic position Genus: Betapartitivirus

Family: Partitiviridae

Geographical distribution DCV2 infection in plants of Anethum graveolens var. hortorum was reported from Germany (Lesker et al. 2013). Symptoms and host(s) The virus-infected plants do not exhibit any external symptoms. Transmission The virus is not mechanically sap-transmissible and is not transmissible by grafting. Virion properties and genome The virions are isometric, non-enveloped, and 30–35 nm in diameter. The genome consists of two double-stranded RNAs. RNA1 consists of 2430 bp (JX971984 = NC_021147) and RNA2 of 2354 bp (JX971985 = NC_021148) (Lesker et al. 2013; Vainio et al. 2018).

References Alhubaishi AA, Walkey DGA, Webb MJW, Bolland CJ, Cook AA (1987) A survey of horticultural plant virus diseases in the Yemen Arab Republic. FAO Plant Prot Bull 35:135–143 Bellardi MG, Rubies-Autonell C (1998) First report of Celery mosaic potyvirus on dill (Anethum graveolens). Plant Dis 82:712 Lesker T, Maiss E (2013) In planta protein interactions of three Alphacryptoviruses and three Betacryptoviruses from white clover, red clover and dill by bimolecular fluorescence complementation analysis. Viruses 5(10):2512–2530 Lesker T, Rabenstein F, Maiss E (2013) Molecular characterization of five betacryptoviruses infecting four clover species and dill. Arch Virol 158(9):1943–1952 Vainio EJ, Chiba S, Ghabrial SA, Maiss E, Roossinck M, Sabanadzovic S, Suzuki N, Xie J, Nibert M, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Partitiviridae. J Gen Virol 99:17–18 Waterhouse PM (1985) Isolation and identification of Carrot red leaf virus from carrot and dill growing in the Australian Capital Territory. Australas Plant Pathol 14:32–34 Wolf P (1972) Virus diseases of dill, Anethum graveolens. Acta Physiol Acad Sci Hung 7(1/3):209–211

Angelica spp. Family: Apiaceae

Medicinal plant

Angelica bushy stunt virus Taxonomic position Genus: Caulimovirus

(AnBSV)

Family: Caulimoviridae

A

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Angelica spp.

Geographical distribution AnBSV infection in plants of Angelica dahurica was reported from South Korea (Lim et al. 2017). Transmission The virus is transmitted by aphid vectors in a semipersistent manner. The virus is also transmitted by mechanical sap inoculation. Virion properties and genome The virions are isometric, not enveloped, 42–45 nm in diameter with no obvious surface structure. The genome is a single molecule of circular dsDNA consisting of 8300 bp (KU508800) with seven putative ORFs (ORFs 1 to 7) containing conserved domains/motifs (Lim et al. 2017).

Angelica virus Y Taxonomic position Genus: Potyvirus

(AVY)

Family: Potyviridae

Geographical distribution AVY infection in wild plants of Angelica genuflexa and A. lucida was reported from Alaska (USA) (Robertson 2007). Symptoms and host(s) The virus-infected angelica plants show leaf mottle symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically saptransmissible. The experimental plant hosts of the virus are Chenopodium amaranticolor and Chenopodium quinoa, Nicotiana benthamiana and N. clevelandii, Anethum graveolens, Apium graveolens var. dulce, Daucus carota, and Petroselinum crispum. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long and 12 nm wide. The genome consists of linear positive-sense single-stranded RNA (Revers and Garcia 2015). Only partial polyprotein sequences of two isolates, 1879 nt (EF488740) and 1888 nt (EF488741), are available (Robertson 2007; Revers and Garcia 2015; Wylie et al. 2017).

Konjac mosaic virus

(KoMV)

Synonyms Japanese hornwort mosaic virus (JHMV) Taxonomic position Genus: Potyvirus

Family: Potyviridae

JHMV-infected plants of Angelica sinensis were reported from China (Zhang et al. 2009). The virusinfected angelica plants exhibit mosaic symptoms. The virus is transmitted by an aphid vector, Myzus

Angelonia angustifolia (Angelonia)

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persicae, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of KoMV, refer to Amorphophallus paeoniifolius.

A References Lim S, Baek D, Igori D, Moon JS (2017) Complete genome sequence of a putative new Caulimovirus which exists as endogenous pararetroviral sequences in Angelica dahurica. Arch Virol 162:3837–3842 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Robertson NL (2007) Identification and characterization of a new virus in the genus Potyvirus from wild populations of Angelica lucida L. and A. genuflexa Nutt., Family: Apiaceae. Arch Virol 152(9):1603–1611 Zhang Y, Wang R, Wang J, Chang J, Zhang X, Chen T, An L, Xu S (2009) A new Potyvirus first isolated and identified from Angelica sinensis. Virus Genes 39:120–125 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Angelonia angustifolia (Angelonia) Family: Plantaginaceae

Ornamental

Alternanthera mosaic virus Taxonomic position Genus: Potexvirus

(AltMV)

Family: Alphaflexiviridae

AltMV infection in plants of Angelonia angustifolia was reported from the USA (Lockhart and Daughtrey 2008). The virus-infected angelonia plants exhibit symptoms of stunting, chlorosis, and light yellow mottle in the foliage, resembling nutrient deficiency. No vector is reported for this virus. The virus is readily sap-transmissible and easily spread by plant contact, vegetative propagation, and contaminated tools and hands. For more details of AltMV, refer to Alternanthera spp.

Angelonia flower break virus

(AFBV)

Synonyms Angelonia flower mottle virus Taxonomic position Genus: Alphacarmovirus

Family: Tombusviridae

Geographical distribution AFBV was isolated from plants of Angelonia angustifolia grown in the USA, Germany, and Israel (Adkins et al. 2006; Winter et al. 2006; Mathews and Dodds 2008). The isolate from Germany was initially named Angelonia flower mottle virus (Winter et al. 2006), which is now recognized as a synonym of AFBV.

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Angraecum spp.

Symptoms and host(s) The virus-infected angelonia plants exhibit mild mottle symptoms on leaves and overall stunting accompanied by flower break. The virus has since been detected in nemesia, verbena, and phlox (Assis Filho et al. 2006; Mathews and Dodds 2008). Transmission The virus is mechanically sap-transmissible, and the experimental host range was limited to Nicotiana spp., Phlox drummondii, and Digitalis purpurea. The virus is not seed-borne in angelonia with high efficiency (Adkins et al. 2006). Virion properties and genome The virions are isometric 30 nm in diameter and have T = 3 icosahedral symmetry. The genome consists of a single molecule of linear positive-sense, single-stranded RNA of 3964 nt (NC_007733 = DQ219415) (Adkins et al. 2006).

References Adkins S, Hammond J, Gera A, Maroon-Lango CJ, Sobolov I, Harness A, Zeidan M, Spiegel S (2006) Biological and molecular characterization of a new Carmovirus isolated from angelonia. Phytopathology 96:460–467 Assis Filho FM, Harness A, Tiffany M (2006) Natural infection of verbena and phlox by a recently described member of the Carmovirus genus. Plant Dis 90:1115 Lockhart BE, Daughtrey ML (2008) First report of Alternanthera mosaic virus infection in Angelonia in the United States. Plant Dis 92:1473 Mathews DM, Dodds JA (2008) First report of Angelonia flower break virus in Nemesia spp. and other ornamental plants in California. Plant Dis 92:651 Winter S, Hamacher A, Engelmann J, Lesemann DE (2006) Angelonia flower mottle, a new disease of Angelonia angustifolia caused by hitherto to unknown Carmovirus. Plant Pathol 55:820

Angraecum spp. Family: Orchidaceae

Ornamental

Cymbidium mosaic virus Taxonomic position Genus: Potexvirus

(CymMV)

Family: Alphaflexiviridae

CymMV infection in plants of Angraecum spp. was detected in Hawaii (Hu et al. 1993). The virusinfected plants exhibit symptoms of elongated, purple-black streaks on the older leaves. Yellow blotches are sometimes visible on the new growth. Older leaves drop prematurely when infection is severe. The virus is mechanially sap-transmissible. The virus also spreads through contact by contaminated cutting tools, equipment, human hands, potting media, or water. No vector transmission is reported. For more details of CymMV, refer to Cymbidium spp.

Annona muricata (Soursop)

141

Orchid fleck dichorhavirus Taxonomic position Genus: Dichorhavirus

(OFV)

Family: Rhabdoviridae

OFV infection in plants of Angraecum spp. was reported from Australia (Kitajima et al. 2001). The virus-infected plants exhibit symptoms of chlorotic and necrotic spots and rings. The virus is transmitted by mite vectors in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of OFV, refer to Cymbidium spp.

References Hu JS, Ferreira S, Wang M, Xu MQ (1993) Detection of Cymbidium mosaic virus, Odontoglossum ringspot virus, Tomato spotted wilt virus, and potyviruses infecting orchids in Hawaii. Plant Dis 77:464–468 Kitajima EW, Kondo H, Mackenzie A, Rezende JAM, Gioria R, Gibbs A, Tamada T (2001) Comparative cytopathology and immunocytochemistry of Japanese, Australian and Brazilian isolates of Orchid fleck virus. J Gen Plant Pathol 67:231–237

Annona muricata (Soursop) Family: Annonaceae

Edible fruit crop

Soursop yellow blotch virus Taxonomic position The virus is a tentative member of the family Rhabdoviridae. Geographical distribution The virus infection in plants of Annona muricata was reported from Ceara, Brazil, by Kitajima et al. (1993). The virus spreads in the South and Central American regions and Brazil (Santos et al. 2007). Symptoms and host(s) The virus-infected soursop plants exhibit symptoms of vein-clearing, yellow blotches on the leaves, leaf distortion, and stunting. Transmission The virus is transmitted by mechanical sap-inoculation to soursop, sweetsop (Annona squamosa), and birida (Rollinia deliciosa). The virus is transmitted by grafting. No seed transmission was observed. Virion properties and genome The virions are rhabdo- or bullet-shaped measuring 250–300 nm in length and 60–70 nm wide (Martins et al. 1999).

A

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Anthocercis spp. (Tailflower)

References Kitajima EW, Martins CRF, Santos AA (1993) Identification of a rhabdovirus in soursop (Annona muricata). Plant Dis 77:276–278 Martins CRF, Lima MI, Barros TSL, Resende RO, Kitajima EW (1999) Further characterization and serological properties of Soursop yellow blotch Rhabdovirus. Fitopatol Bras 24:410–415 Santos AA d, Cardoso JE, Viana FMP, Vidal JC, Souza RNM (2007) Effect of the Soursop yellow blotch virus on the growth and yield of soursop diseased plants. Summa Phytopathol 33:90–92

Anthocercis spp. (Tailflower) Family: Solanaceae

Weed host

Pelargonium zonate spot virus Taxonomic position Genus: Anulavirus

(PZSV)

Family: Bromoviridae

PZSV infection in plants of Anthocercis ilicifolia was reported from Australia (Li et al. 2016). The virus-infected tailflower plants exhibit chlorosis and necrosis symptoms. Thrips are involved in transmitting this virus by carrying the virus-infected pollen grains on their bodies and introducing the virus into the susceptible hosts while feeding. The virus is mechanically sap-transmissible to a wide range of test plants. For more details of PZSV, refer to Pelargonium spp.

Yellow tailflower mild mottle virus Taxonomic position Genus: Tobamovirus

(YTMMV)

Family: Virgaviridae

Geographical distribution YTMMV infection in plants of Anthocercis spp. was first reported from Australia (Wylie et al. 2014; Li et al. 2016). Symptoms and host(s) The virus-infected tailflower plants exhibit symptoms of mild mottling and chlorosis on the leaves. Transmission No vector is known for this virus. The virus is mechanically sap-transmissible to seedlings of Nicotiana glutinosa, N. benthamiana, N. umbratica, Chenopodium quinoa, and C. amaranticolor and produces systemic symptoms, sometimes severe on Nicotiana plants and local lesions on Chenopodium. The virus is also transmissible by contact between plants. Virion properties and genome The virions have rigid rod-shaped morphology measuring 240  14 nm. The genome is a monopartite, linear, positive-sense ssRNA of 6379 nt (KF495564 = NC_022801). The 30 -terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G5’pppG) (Zaitlin 2011; Wylie et al. 2014; Adams et al. 2017).

Anthoxanthum odoratum (Sweet vernalgrass)

143

References Adams MJ, Adkins S, Bragard C, Gilmer D, Li D, MacFarlane SA, Wong S-M, Melcher U, Ratti C, Ryu KH, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Virgaviridae. J Gen Virol 98:1999–2000 Li H, Zhang C, Luo H, Jones MGK, Sivasithamparam K, Koh S-H, Ong JWL, Wylie SJ (2016) Yellow tailflower mild mottle virus and Pelargonium zonate spot virus co-infect a wild plant of red-striped tailflower in Australia. Plant Pathol 65:503–509 Wylie SJ, Li H, Jones MG (2014) Yellow tailflower mild mottle virus: a new Tobamovirus described from Anthocercis littorea (Solanaceae) in Western Australia. Arch Virol 159(4):791–795 Zaitlin M (2011) Tobamovirus. In: Tidona C, Darai G (eds) The Springer index of viruses. Springer, New York. https://doi. org/10.1007/978-0-387-95919-1

Anthoxanthum odoratum (Sweet vernalgrass) Family: Poaceae

Weed host

Anthoxanthum latent blanching virus Taxonomic position Genus: Hordeivirus

(ALBV)

Family: Virgaviridae

Geographical distribution ALBV infection in plants of Anthoxanthum odoratum was first reported from Wales, UK (Catherall and Chamberlain 1980). The virus spreads in the UK (Edwards et al. 1989). Symptoms and host(s) The virus-infected sweet vernal grass plants express inconspicuous chlorotic stripes in leaves and are sometimes symptomless. Transmission The virus is mechanically sap-transmissible to graminaceous hosts. There is no known vector for this virus. Virion properties and genome The virions are rod-shaped, straight, with modal length of 20  110–150 nm diameter (Edwards et al. 1989). The genome consists of three positive-sense single-stranded RNAs and a “triple gene block” set of cell-to-cell movement proteins (Jackson et al. 2011; Adams et al. 2017).

References Adams MJ, Adkins S, Bragard C, Gilmer D, Li D, MacFarlane SA, Wong S-M, Melcher U, Ratti C, Ryu KH, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Virgaviridae. J Gen Virol 98:1999–2000 Catherall PL, Chamberlain JA (1980) A new virus disease of sweet vernal grass (Anthoxanthum odoratum). In: Proceedings of the 3rd conference on virus diseases of Gramineae in Europe. Rothamsted, 28–30 May, pp 83–85 Edwards ML, Kelley SE, Arnold MK, Cooper JL (1989) Properties of a Hordeivirus from Anthoxanthum odoratum. Plant Pathol 38:209–218 Jackson AO, Bragg J, Lim HS, Ganesan U (2011) Hordeivirus. In: Tidona C, Darai G (eds) The Springer index of viruses. Springer, New York. https://doi.org/10.1007/978-0-387-95919-1

A

144

Anthriscus cerefolium (Chervil)

Anthriscus cerefolium (Chervil) Family: Apiaceae

Medicinal

Carrot red leaf virus

(CtRLV)

Taxonomic position Genus: Polerovirus

Family: Luteoviridae

CtRLV infection in plants of Anthriscus cerefolium was reported from Scotland (Murant and Roberts 1979). The virus is not mechanically sap-transmissible but is transmitted by the aphid vectors in a circulative, non-propagative manner. For more details of CtRLV, refer to Daucus carota.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Anthriscus cerefolium was reported from Austria (Grausgruber-Groger 2012). The virus-infected chervil plants exhibit necrotic spot symptoms on the leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

References Grausgruber-Groger S (2012) First report of Impatiens necrotic spot virus on Ocimum basilicum, Eruca sativa and Anthriscus cerefolium in Austria. New Dis Rep 26:12 Murant AF, Roberts IM (1979) Virus-like particles in phloem tissue of chervil (Anthriscus cerefolium) infected with Carrot red leaf virus. Ann Appl Biol 92:343–346

Anthriscus sylvestris (Cow parsley) Family: Apiaceae

Weed host

Anthriscus yellows virus Taxonomic position Genus: Waikavirus

(AYV)

Family: Secoviridae

Geographical distribution AYV infection in plants of Anthriscus sylvestris was first reported from the UK by Murant and Goold (1968). The virus spreads in the Eurasian region and the UK (Hemida et al. 1989).

Anthurium andraeanum (Flamingo flower)

145

Symptoms and host(s) The virus-infected cow parsley plants do not exhibit conspicuous symptoms.

A Transmission The virus is transmitted by an aphid vector, Cavariella aegopodii in a semi-persistent manner. The virus can help the vector transmission of another virus (Parsnip yellow fleck sequivirus) (Murant and Goold 1968; Elnagar and Murant 1976). The virus is not transmitted by mechanical inoculation. Virion properties and genome The virions are non-enveloped 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is a monopartite linear positive-sense, single-stranded RNA (Hemida et al. 1989; Sanfacon et al. 2009; Hibino 2011; Sanfacon 2015; Thompson et al. 2017).

References Elnagar S, Murant AF (1976) Relations of the semi-persistent viruses, Parsnip yellow fleck and Anthriscus yellows, with their vector, Cavariella aegopodii. Ann Appl Biol 84:153–167 Hemida SK, Murant AF, Duncan GH (1989) Purification and some particle properties of Anthriscus yellows virus, a phloem limited semi-persistent aphid-borne virus. Ann Appl Biol 114:71–86 Hibino H (2011) Waikavirus. Sequiviridae. In: The Springer index of viruses. Springer, New York, pp 1775–1779. https:// doi.org/10.1007/978-0-387-95919-1_289 Murant AF, Goold RA (1968) Purification, properties and transmission of Parsnip yellow fleck, a semi-persistent, aphidborne virus. Ann Appl Biol 62:123–137 Sanfacon H (2015) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. In: eLS. Wiley, Chichester. http://www.els.net https://doi.org/10.1002/9780470015902.a0000764.pub3 Sanfacon H, Wellink J, Le Gall O, Karasev A, van der Vlugt R, Wetzel T (2009) Secoviridae: a proposed family of plant viruses within the order Picornavirales that combines the families Sequiviridae and Comoviridae, the unassigned genera Cheravirus and Sadwavirus, and the proposed genus Torradovirus. Arch Virol 154(5):899–907 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J Gen Virol 98:529–531

Anthurium andraeanum (Flamingo flower) Family: Araceae

Ornamental

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Anthurium andraeanum was reported from Brazil and Iran (Ghotbi and Nazerian 2010; Miura et al. 2013). The virus-infected flamingo flower plants exhibited intensive mosaic and leaf deformation symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a number of hosts. For more details of CMV, refer to Cucumis sativus.

146

Dasheen mosaic virus Taxonomic position Genus: Potyvirus

Anthurium andraeanum (Flamingo flower)

(DsMV)

Family: Potyviridae

DsMV infection in plants of Anthurium andraeanum was reported from Brazil, and the USA (Elliott et al. 1997; Lima et al. 2004; Alexandre et al. 2005; Lima and Nascimento 2012; Zavareh et al. 2013; Ghotbi et al. 2016). The virus-infected flamingo flower plants exhibited mosaic and chlorotic stripes along the foliar veins and reduced plant growth. Color degradation in the flowers and bractless deformations, with serious consequences in the appearance and quality of the flowers, were also recorded (Rivas et al. 1997). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of DsMV, refer to Colocasia esculenta.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

Localized infection of Anthurium andraeanum was detected in a greenhouse in the Czech Republic (Mertelik et al. 2002) and in Anthurium spp. in Iran (Ghotbi and Shahraeen 2012; Ghotbi 2013). The virus-infected flamingo flower plants exhibit symptoms of necrotic leaf spots, chlorosis, and stunting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Anthurium andraeanum was reported from the USA, the Netherlands, Portugal, and Iran (Verhoeven and Roenhorst 1992; Louro 1996; Uchida et al. 1999; Ghotbi and Shahraeen 2012). The virus-infected flamingo flower plants show that initial symptoms of 5–10 mm chlorotic patches. These were subsequently developing shallow brown pitting within the chlorotic area and brown necrotic edges to the patches and expanding to 20–30 mm in diameter. The leaves and spathes may develop water-soaked spots resembling bacterial blight, but lacking bacterial ooze; young shoots and sections of the plant may be killed by infection. Necrotic lesions on flowers are noticed. The virus is transmitted by thrips vectors Frankliniella occidentalis, F. schultzei, and Thrips tabaci in a persistent-propagative manner (Uchida et al. 1999). The virus is also transmissible by mechanical sapinoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Alexandre MAV, Rivas EB, Tozetto ARP, Duarte LML (2005) An annotated list on the natural occurrence of viruses in ornamental plants in Brazil. Instituto Biologico, Sao Paulo, p 54

Antirrhinum spp. (Snapdragon)

147

Elliott MS, Zettler FW, Brown LG (1997) Dasheen mosaic potyvirus of edible and ornamental aroids. Plant Pathology Circular No. 384, Fla Dept and Consumer Services, July/August 1997 Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iran J Plant Path 49:41 Ghotbi T, Nazerian I (2010) Report on incidence of Cucumber mosaic virus (CMV) on ornamental Anthurium in Iran. In: 19th Iranian plant protection congress, Tehran. 722 Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses affecting propagated ornamentals in northern Iran. Int Res J Microbiol 3:373–381 Ghotbi T, Shahraeen N, Alaee A (2016) Occurrence of Dasheen mosaic virus on Anthurium ornamental plant in Iran. In: National conference on applied research in agriculture and natural resources. Agriculture and Natural Resources Engineering Disciplinary Organization. Karaj, Iran. Lima JAA, Nascimento AKQ (2012) Identification of Dasheen mosaic virus in Anthurium andraeanum by immune precipitation polymerase chain reaction in Brazil. Bioforsk Fokus 7(9):24 Lima RCA, Lima JAA, Rubens Aguiar J (2004) Serological identification of Dasheen mosaic virus in Anthurium sp. in the State of Ceara. Fitopatol Brazil 29:105 Louro D (1996) Detection and identification of Tomato spotted wilt virus and Impatiens necrotic spot virus in Portugal. Acta Hortic 431:99–105 Mertelik J, Mokra V, Gotzova B, Gabrielova S (2002) Occurrence and identification of Impatiens necrotic spot tospovirus in the Czech Republic. Acta Hortic 568:79–83 Miura NS, Beriam LOS, Rivas EB (2013) Detection of Cucumber mosaic virus in commercial anthurium crops and genotypes evaluation. Hortic Bras 31:322–327 Rivas EB, Duarte LML, Alexandre MAV, Galleti SR (1997) Dasheen mosaic virus in Anthurium species. Virus Rev Res 2:192–193 Uchida JY, Ogata D, Nagata N (1999) Tomato spotted wilt virus on anthurium. University of Hawaii Cooperative Extension Service, Honolulu, p 1 (Plant Disease; PD-17) Verhoeven JTJ, Roenhorst JW (1992) Tomato spotted wilt virus: ecological aspects in ornamental crops in the Netherlands from 1989 up to 1991. Acta Hortic 377:175–182 Zavareh N, Maleki M, Ghotbi T (2013) Serological and molecular detection of Cucumber mosaic virus from two main commercial anthurium cultivars in Iran. Ann Biol Res 4(4):120–125

Antirrhinum spp. (Snapdragon) Family: Plantaginaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Antirrhinum majus was reported from Argentina (Arneodo et al. 2005). The virus-infected snapdragon plants exhibit mosaic symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(INSV)

A

148

Antirrhinum spp. (Snapdragon)

Infection of snapdragon (Antirrhinum majus) by INSV has been reported from many countries including the USA and Italy (Daughtrey 1996; Daughtrey et al. 1997; Vaira et al. 1993), causing symptoms including stem discoloration, wilt, and white leaf spotting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato yellow ring virus

(TYRV)

Synonyms Tomato Varamin virus (ToVV)

Taxonomic position TYRV is a tentative member of the genus Orthotospovirus and family Tospoviridae Plants of Antirrhinum spp. were found to be infected with ToVV in Iran (Ghotbi et al. 2005). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sapinoculation. For more details of TYRV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Antirrhinum majus was reported from the USA (Cho et al. 1987; Daughtrey 1996; Dobhal et al. 2014). The virus-infected snapdragon plants exhibit symptoms of yellow leaf spots or blotches. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is not transmissible by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Arneodo JD, Guzman FA, Nome CF, Conci LR (2005) Occurrence of Cucumber mosaic virus on Antirrhinum majus in Argentina. Fitopatol Bras 30:90 Cho JJ, Mau RFL, Mitchell WC, Gonsalves D, Yudin LS (1987) Host list of plants susceptible to Tomato spotted wilt virus (TSWV). University of Hawaii, Honolulu, p 10 (Research Extension Series; RES-078) Daughtrey ML (1996) Detection and identification of tospoviruses in greenhouses. Acta Hortic 431:90–98 Daughtrey ML, Jones RK, Moyer JW, Daub ME, Baker JR (1997) Tospoviruses strike the greenhouse industry: INSV has become a major pathogen on flower crops. Plant Dis 81(11):1220–1230 Dobhal S, Arif M, Olson J, Mendoza-Yerbafria A, Aguilar-Moreno S, Perez-Garcia M, Ochoa-Corona FM (2014) Sensitive detection and discrimination method for studying multiple infections of five major plant viruses infecting ornamental plants in nursery environments. Ann Appl Biol 166:286–296 Ghotbi T, Sharaeen N, Winter S (2005) Occurrence of tospoviruses in ornamental and weed species in Markazi and Tehran provinces in Iran. Plant Dis 89:425–429 Vaira AM, Roggero P, Luisoni E, Masenga V, Milne RG, Lisa V (1993) Characterization of two tospoviruses in Italy: tomato spotted wilt and impatiens necrotic spot. Plant Pathol 42:530–542

Aphelandra squarrosa (Zebra plant)

149

Apera spica-venti (Silky bent grass) Family: Poaceae

Wheat dwarf virus Taxonomic position Genus: Mastrevirus

A

Fodder

(WDV)

Family: Geminiviridae

WDV infection in plants of Apera spica-venti was reported from the Czech Republic (Vacke and Cibulka 1999). The virus-infected Silky bent grass plants exhibit symptoms of dwarfing, leaf yellowing, and inflorescence reduction. The virus is transmitted by a leafhopper vector Psammotettix alienus in a persistent, circulative, and non-propagative manner. The virus is not transmissible by mechanical inoculation, nor transmissible by contact between plants. The virus is not transmitted by seed and pollen. For more details of WDV, refer to Triticum aestivum.

Reference Vacke J, Cibulka R (1999) Silky bent grass (Apera spica-venti [L.] Beauv.) – a new host and reservoir of Wheat dwarf virus. Plant Protect Sci 35:47–50

Aphelandra squarrosa (Zebra plant) Family: Acanthaceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Aphelandra squarrosa was reported from Portugal (Louro 1996). The virusinfected zebra plants exhibit symptoms such as veinal necrosis, distortion of young growth, stunting, and wilting. These symptoms can vary with the stage of growth of the host and with a variety of cultural conditions. The virus is transmitted by a thrips vector Frankliniella occidentalis in a persistentpropagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

150

Apios americana (Wild groundnut)

TSWV infection in plants of Aphelandra squarrosa was reported from Portugal and the USA (Halliwell and Barnes 1987; Louro 1996). The virus-infected zebra plants exhibit progressive light brown necrosis of the major veins of semi-mature terminal leaves. The discoloration frequently progressed to involve large intervenial areas resulting in extensive leaf necrosis. Symptomatic leaves were distorted and epinastic and had a tendency to abscise prematurely. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Halliwell RS, Barnes LW (1987) Tomato spotted wilt virus infection of commercial Aphelandra sp. J Envir Hortic 5:120–121 Louro D (1996) Detection and identification of Tomato spotted wilt virus and Impatiens necrotic spot virus in Portugal. Acta Hortic 431:99–105

Apios americana (Wild groundnut) Family: Fabaceae

Tuber crop

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Apios americana was reported from Louisiana (USA) (Valverde et al. 1990). The virus-infected wild groundnut plants exhibit foliar mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Desmodium yellow mottle virus Taxonomic position Genus: Tymovirus

(DYMoV)

Family: Tymoviridae

DYMoV infection in plants of Apios americana was reported from Louisiana (USA) (Valverde et al. 1990). The virus-infected wild groundnut plants exhibit mild yellow mottle symptoms. The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of DYMoV, refer to Desmodium spp.

Reference Valverde RA, Provvidenti R, Clark CA (1990) Cucumber mosaic virus and Desmodium yellow mottle virus infections in wild groundnut (Apios americana). Plant Dis 74:151–153

Apium graveolens (Celery)

151

Apium graveolens (Celery) Family: Apiaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Leafy vegetable

(AMV)

Family: Bromoviridae

AMV infection in plants of Apium graveolens was reported from Israel (Marco 1975). The virusinfected celery plants exhibit prominent stunting and a bright yellow mottle of a calico-like mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Apium virus Y

(ApVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution ApVY occurs in plants of Apium graveolens from the USA and New Zealand (Tang et al. 2007; Eastwell et al. 2008; Tian et al. 2008; Xu et al. 2011a; Koike et al. 2012). Symptoms and host(s) The virus-infected celery plants older leaves generally show yellow or brown line patterns, yellow blotches, and brown lesions and in some cases distorted and twisted leaflets. Younger leaves may show a faint mosaic or mottling. The virus-infected celery petioles could exhibit dark brown, sunken, elongated lesions which reduce the marketability. However, for some cultivars, the celery plants having symptomatic foliage will not have any petiole symptoms. Transmission This virus is vectored by an aphid, Myzus persicae, in a non-persistent manner. The virus is also mechanically sap-transmissible. The experimental host range of the virus includes 13 plant species in the families Apiaceae, Chenopodiaceae, and Solanaceae. No seed transmission has been reported. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long and 11–13 nm wide. The genome is a single molecule of linear, positive-sense single-stranded RNA of 9917 nt (HM363516 = NC_014905) (Tian et al. 2008; Xu et al. 2011a; Revers and Garcia 2015; Wylie et al. 2017).

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

A

152

Apium graveolens (Celery)

ArMV infection in plants of Apium graveolens was reported from New Zealand (Mossop et al. 1983). The virus-infected celery plants exhibit symptoms of chlorosis, vein necrosis, leaf distortion, and marked reduction in vigor. The virus is transmitted by nematode vector in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Celery mosaic virus

(CeMV)

Synonyms Western celery mosaic virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution CeMV infection in plants of Apium graveolens was reported from Britain, France, Germany, Yugoslavia, Poland, the Netherlands, Italy, Japan, Argentina, Canada, France, Czechoslovakia, Venezuela, New Zealand, Australia, the UK, and the USA (Zitter 1970; Kemp and Frowd 1975; Alberts et al. 1989; Traicevski et al. 1999; Raid and Zitter 2002; Latham and Jones 2001, 2003; Fernandez et al. 2006; Paduch-Cichal and Sala-Rejczak 2010; Amal et al. 2011). Symptoms and host(s) Symptoms incited by CeMV in infected celery plants include yellowing of foliage, mosaic and/or mottling patterns on leaves, vein-clearing, and curled, crinkled, or otherwise distorted foliage. On older leaves necrotic leaf spots may develop. Plants are sometimes stunted. Outer petioles grow horizontally, giving the plant a flattened appearance. Celery plants that become infected in the early to middle part of their growing cycle are unmarketable, while plants that become infected late may be marketable but with a reduced shelf life. Certain strains of the virus may cause more severe symptoms than other strains. The natural hosts of this virus are limited to specie in the Umbelliferae. Economically important hosts include celery, carrot, parsnip, parsley, dill, caraway, and hemlock weed host. Transmission The virus is transmitted by a number of aphid species including Myzus persicae, M. circumflexus, M. convolvuli, Aphis middletonii, A. ferruginea-striata, A. apigraveolens, A. apii, A. gossypii, A. rumicis, Cavariella capreae, C. aegopodii, C. pastinacae, and Rhopalosiphum melliferum in a non-persistent manner. The virus is also mechanically sap-transmissible and has a limited host range of fewer than three plant families. The virus is not transmissible by contact between plants. The virus is not reported to be seed-borne. Virion properties and genome The virions are non-enveloped, flexuous filaments, 780 nm long and 15 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 9999 nt (HQ676607 = NC_015393) (Shepard and Grogan 1967; Xu et al. 2011b; Revers and Garcia 2015; Wylie et al. 2017).

Apium graveolens (Celery)

153

Celery yellow mosaic virus

(CeYMV)

Taxonomic position CeYMV is a tentative member of the genus Potyvirus and family Potyviridae. Geographical distribution CeYMV infection in plants of Apium graveolens was reported from Brazil and Egypt (Kitajima and Costa 1968; Oliveira and Kitajima 1981; Abo El-Ela et al. 2005). Symptoms and host(s) The virus-infected celery plants exhibit conspicuous leaf chlorosis symptoms. Transmission The virus is transmitted by an aphid vector, Myzus persicae, in a non-persistent manner, and the virus is also mechanically sap-transmissible. Virion properties and genome The virions are flexuous filaments with a clear modal length of 750 nm. The genome is a single molecule of positive-sense single-stranded RNA.

Chickpea chlorotic stunt virus Taxonomic position Genus: Polerovirus

(CpCSV)

Family: Luteoviridae

CpCSV infection in plants of Apium graveolens was reported from Syria (Asaad et al. 2009). The virusinfected celery plants exhibit symptoms of yellowing, reddening, and/or stunting. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not mechanically saptransmissible. For more details of CpCSV, refer to Cicer arietinum.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMVoccurs in Apium graveolens wherever the crop is grown (Zitter 1970; Bruckart and Lorbeer 1975, 1976; Marco 1975; Pemberton and Frost 1986). CMV infection of celery is referred to as southern bean mosaic virus in the older literature following its description in southern production areas. The virusinfected celery leaves will develop vein-clearing and mosaic, and later these same leaves may show yellowing and veinal necrosis. The petioles of these plants show elongated, brown to translucent, sunken beige-colored lesions, making the celery stalk unmarketable. Symptoms may also be transient, such that later in the season, the symptoms may be limited to a dull cast, while under cool growing conditions, the obvious symptoms may be muted. The virus is transmitted by more than 60 aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

A

154

Apium graveolens (Celery)

Parsnip yellow fleck virus

(PYFV)

Synonyms Celery yellow net virus Taxonomic position Genus: Sequivirus

Family: Secoviridae

PYFV was first reported in plants of Apium graveolens by Hollings (1964) from the UK. The virus spreads in Europe. The virus-infected celery plants exhibit yellow flecking, vein-banding, and leaf malformation symptoms. The virus is transmitted by aphid vectors in a semi-persistent manner, requires for vector transmission a helper virus (anthriscus yellows waikavirus). The virus is transmitted by mechanical sap-inoculation to less than three family members. For more details of PYFV, refer to Pastinaca sativa.

Strawberry latent ringspot virus Taxonomic position Genus: Unassigned

(SLRSV)

Family: Secoviridae

SLRSV infection in plants of Apium graveolens was reported from the USA and England (Walkey and Mitchell 1969). The young leaves of virus-infected celery plants are narrow with distorted and shortened petioles. The virus is transmitted by nematode vectors (Xiphinema spp.) and also by mechanical sap-inoculation to a very wide host range. The virus is transmitted by grafting. The virus is seed-transmitted in celery plants to the extent of 98–100% (Walkey and Whittingham-Jones 1970). For more details of SLRSV, refer to Fragaria spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Apium graveolens was reported from China, Argentina, and New Zealand (Mossop et al. 1983; Gracia and Feldman 1989; Dellavalle et al. 1999; Li et al. 2015). The virusinfected celery plants exhibit symptoms of yellow spots that turn brown on the older leaves. Large sections of stalks turn brown and eventually die; plants may be severely stunted. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Abo El-Ela AA, Amer MA, Abo El-Abbas F (2005) Celery yellow mosaic potyvirus affecting umberlliferae plants in Egypt. Egypt J Virol 2(1):269–282 Alberts E, Francki RIB, Dietzgen RG (1989) An epidemic of Celery mosaic virus in South Australian celery. Aust J Agric Res 40:1027–1036

Apium graveolens (Celery)

155

Amal AA, Zein NS, Khatab AH (2011) Characterization of Celery mosaic virus isolated from some Apiaceae plants. Int J Virol 8(2):214–223 Asaad NY, Kumari SG, Haj-Kassem AA, Shalaby A-BA, Al-Shaabi S, Malhotra RS (2009) Detection and characterization of Chickpea chlorotic stunt virus in Syria. J Phytopathol 157:756–761 Bruckart WL, Lorbeer JW (1975) Recent occurrences of cucumber mosaic, lettuce mosaic and broadbean wilt viruses in lettuce and celery fields in New York. Plant Dis Reptr 59:203–206 Bruckart WL, Lorbeer JW (1976) Cucumber mosaic virus in weed hosts near commercial fields of lettuce and celery. Phytopathology 66:253–259 Dellavalle G, Ciuffo M, Roggero P, Lisa V, Minuto A, Minuto G, Rapetti S (1999) Tospoviruses in Delphinium sp., gazania, marguerite, celery, Tragopogon porrifolius and Solanum rantonnetti in Liguria (northern Italy). Informatore Fitopatologico 49:63–64 Eastwell KC, Glass JR, Seymour LM, Druffel KJ (2008) First report of infection of poison hemlock and celery by Apium virus Y in Washington State. Plant Dis 92:1710 Fernandez T, Carballo O, Zambrano K, Romano M, Marys E (2006) First report of Celery mosaic virus infecting celery in Venezuela. Plant Dis 90:1111 Gracia O, Feldman JM (1989) First report of Tomato spotted wilt virus on celery and three weed species in Argentina. Plant Dis 73:859 Hollings M (1964) Some properties of five viruses of celery (Apium graveolens L.) in Britain. J Hortic Sci 39:130–141 Kemp WG, Frowd A (1975) The isolation and identification of Celery mosaic virus in Ontario. Plant Dis Reptr 59:50–53 Kitajima EW, Costa AS (1968) Celery yellow mosaic virus. Bragantia 27:7–12 Koike ST, Liu H-Y, Sears J, Tian T, Daugovish O, Dara S (2012) Distribution, cultivar susceptibility and epidemiology of Apium virus Y on celery in coastal California. Plant Dis 96:612–617 Latham L, Jones RAC (2001) ‘Celery mosaic virus.’ Agriculture Western Australia Fact Sheet No. 15 Latham LJ, Jones RAC (2003) Incidence of Celery mosaic virus in celery crops in south-west Australia and its management using a ‘celery-free period’. Australas Plant Pathol 32:527–531 Li YY, Xiao L, Tan GL, Fu XP, Li RH, Li F (2015) First report of Tomato spotted wilt virus on celery in China. Plant Dis 99:734 Marco S (1975) Occurrence and incidence of Cucumber mosaic virus and Alfalfa mosaic virus on celery in Israel. Plant Dis Reptr 59:915–917 Mossop DW, Fry PR, Young BR (1983) New plant disease records in New Zealand: Arabis mosaic virus in celery, lettuce and Chinese cabbage; Tomato spotted wilt virus in celery. NZ J Agric Res 26:257–259 Oliveira ML, Kitajima EW (1981) Biological properties of Celery yellow mosaic virus. Fitopatol Bras 6:35–46 Paduch-Cichal E, Sala-Rejczak K (2010) Celery mosaic in celery in Poland. J Plant Pathol 92:S4.113 Pemberton AW, Frost RR (1986) Virus diseases of celery in England. Ann Appl Biol 108:319–331 Raid RN, Zitter TA (2002) Celery mosaic. In: Davis M, Raid R (eds) Compendium of umbelliferous diseases. American Phytopathological Society, Minnesota, pp 53–54 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Shepard JF, Grogan RG (1967) Partial purification, properties and serology of Western celery mosaic virus. Phytopathology 57:1104–1110 Tang J, Clover GRG, Alexander BJR (2007) First report of Apium virus Y in celery in New Zealand. Plant Dis 91:1682 Tian T, Liu H-Y, Koike ST (2008) First report of Apium Virus Y on cilantro, celery, and parsley in California. Plant Dis 92:1254 Traicevski V, Schreurs T, Rodoni B, Moran J (1999) Celery mosaic potyvirus occurring naturally in cultivated Apiaceae in Victoria, Australia. Australas Plant Pathol 28:92 Walkey DGA, Mitchell L (1969) Studies on a ‘strap-leaf’ disease of celery caused by Strawberry latent ringspot virus. Plant Pathol 18:167–172 Walkey DGA, Whittingham-Jones SG (1970) Seed-transmission of Strawberry latent ringspot virus in celery (Apium graveolens var.dulce). Plant Dis Reptr 54:802–803 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Xu D, Liu H-Y, Koike ST, Li F, Li R (2011a) Biological characterization and complete genomic sequence of Apium virus Y infecting celery. Virus Res 155:76–82 Xu D, Liu HY, Li F, Li R (2011b) Complete genome sequence of Celery mosaic virus and its relationship to other members of the genus Potyvirus. Arch Virol 156(5):917–920 Zitter TA (1970) Cucumber mosaic and western celery mosaic-two aphid-transmitted virus diseases of Florida celery. Proc Florida State Hortic Soc 83:188–191

A

156

Aporcactus flagelliformis (Rattail cactus)

Aporcactus flagelliformis (Rattail cactus) Family: Cactaceae

Ornamental

Rattail cactus necrosis-associated virus Taxonomic position Genus: Tobamovirus

(RCNaV)

Family: Virgaviridae

Geographical distribution RCNaV infection in plants of Aporcactus flagelliformis was reported from Korea (Kim et al. 2012). Symptoms and host(s) The virus-infected rattail cactus plants exhibit necrosis symptoms.

Transmission The virus is mechanically sap-transmissible and produced necrotic local lesions on Gomphrena globosa, Chenopodium amaranticolor, C. quinoa, Datura stramonium, Nicotiana benthamiana, and N. rustica. Systemic necrosis symptoms were produced on Nicotiana benthamiana. The virus is also transmissible by contact between plants. There is no known vector for this virus.

Virion properties and genome The virions are rod shaped and 18 nm in diameter and have a predominant length of 300–310 nm. The genome is a monopartite, linear, positive-sense ssRNA of 6,506 nt and contained four open reading frames coding for proteins of M(r) 128 kDa (3,441 nt), 185 kDa (4,929 nt), 55 kDa (1,452 nt), 36 kDa (1,005 nt), and 19 kDa (513 nt) from the 50 to 30 end, respectively (JF729471 = NC_016442) (Kim et al. 2012). The 30 terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G50 pppG).

Reference Adams MJ, Adkins S, Bragard C, Gilmer D, Li D, MacFarlane SA, Wong S-M, Melcher U, Ratti C, Ryu KH, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Virgaviridae. J Gen Virol 98:1999–2000 Kim NR, Hong JS, Song YS, Chung BN, Park JW, Ryu KH (2012) The complete genome sequence of a member of a new species of tobamovirus (Rattail cactus necrosis-associated virus) isolated from Aporcactus flagelliformis. Arch Virol 157:185–187 Zaitlin M (2011) Tobamovirus. In: Tidona C, Darai G (eds) The Springer index of viruses. Springer, New York. https://doi. org/10.1007/978-0-387-95919-1

Aquilegia vulgaris (Columbine)

157

Aquilegia vulgaris (Columbine) Family: Ranunculaceae

A

Medicinal

Aquilegia necrotic mosaic virus

(ANMV)

Taxonomic position ANMV is a tentative member of the genus Caulimovirus and family Caulimoviridae. Geographical distribution ANMV was first reported in plants of Aquilegia spp. from Japan by Y. Doi (in Hull et al. 1989). The virus spreads in the Eastern Asian region including Japan, as well as in Colombia, and East Africa. Symptoms and host(s) The virus-infected columbine plants exhibit leaf chlorosis and necrosis symptoms. Virion properties and genome The virions are isometric, non-enveloped, 50 nm in diameter, and rounded in profile. The genome consists of double-stranded circular DNA.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Aquilegia spp. was reported from Ohio, USA (Fisher et al. 1997). The virusinfected columbine plants exhibit symptoms of mosaic, chlorosis, leaf curl, and stunting. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sapinoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

References Fisher J, Sanchez-Cuevas M-C, Nameth S, Woods V (1997) First report of Cucumber mosaic virus in Eryngium amethystinum, Canna spp., and Aquilegia hybrids in Ohio. Plant Dis 81:1331 Hull R, Brown F, Payne CC (1989) Virology: directory and dictionary of animal, bacterial and plant viruses. Macmillan Press, London, p 11

158

Arabidopsis spp.

Arabidopsis spp. Family: Brassicaceae

Weed host

Arabidopsis halleri partitivirus 1 Taxonomic position The virus is a tentative member of the genus Alphapartitivirus and family Partitiviridae. Geographical distribution The virus infection in plants of Arabidopsis halleri was reported from Japan (Kamitani et al. 2016). Transmission The virus is transmitted through pollen and seed. Virion properties and genome The virions are isometric, 25–50 nm in diameter. The genome consists of double-stranded RNA of two parts. RNA1 consists of 1959 nt (LC151461) and RNA2 is 1763 nt (LC151462) (Vainio et al. 2018).

Cauliflower mosaic virus Taxonomic position Genus: Caulimovirus

(CaMV)

Family: Caulimoviridae

CaMV infection in plants of Arabidopsis thaliana was reported from France, and Oklahoma, USA (Balazs and Lebeurier 1981; Melcher 1989). The virus-infected thale cress plants exhibit vein-clearing, chlorotic spotting, and stunting symptoms. The virus is transmitted by aphid vectors in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of CaMV, refer to Brassica oleracea var. botrytis.

Melandrium yellow fleck virus Taxonomic position Genus: Bromovirus

(MYFV)

Family: Bromoviridae

MYFV infection in plants of Arabidopsis thaliana was reported from Japan (Narabayashi et al. 2009). The virus is transmitted by beetle vectors and also by mechanical sap-inoculation. For more details of MYFV, refer to Silene spp.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

Arabidopsis spp.

159

TMV infection in plants of Arabidopsis thaliana was reported from Georgia, USA (de Assis Filho and Sherwood 2000). The virus-infected thale cress plants exhibit flower abortion symptoms. There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. A high percentage of seed transmission is reported in A. thaliana (de Assis Filho and Sherwood 2000). For more details of TMV, refer to Nicotiana tabacum.

Turnip yellow mosaic virus Taxonomic position Genus: Tymovirus

(TYMV)

Family: Tymoviridae

TYMV infection in plants of Arabidopsis thaliana was reported from Georgia, USA (de Assis Filho and Sherwood 2000). The most common visible symptoms in A. thaliana infected with TYMV were mosaic, stunting, delayed flowering, flower abortion, and reduction in the size of leaves, flowers, and fruits. The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sap-inoculation. The virus is seed-transmitted in Arbidopsis thaliana plants to the extent of 72.6% (de Assis Filho and Sherwood 2000). Thus, embryo invasion is necessary for seed transmission of TYMV in A. thaliana. The virus is being carried by pollen to the seed. For more details of TYMV, refer to Brassica rapa.

Youcai mosaic virus

(YoMV)

Synonyms Oilseed rape mosaic virus Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

YoMV infection in plants of Arabidopsis thaliana was reported from Spain (Aguilar et al. 1996). The virus-infected thale cress plants exhibit mosaic symptoms. There is no known vector for this virus. The virus is mechanically sap-transmissible and also through contact between plants. For more details of YoMV, refer to Brassica napus.

References Aguilar I, Sanchez F, Martin AM, Martinez-Herrera D, Ponz F (1996) Nucleotide sequence of Chinese rape mosaic virus (Oilseed rape mosaic virus), a crucifer Tobamovirus infectious on Arabidopsis thaliana. Plant Mol Biol 30:191–197 Balazs E, Lebeurier G (1981) Arabidopsis is a host of Cauliflower mosaic virus. Arabidopsis Inf Serv 18:130–134 de Assis Filho FM, Sherwood JL (2000) Evaluation of seed transmission of Turnip yellow mosaic virus and Tobacco mosaic virus in Arabidopsis thaliana. Phytopathology 90:1233–1238 Kamitani M, Nagano AJ, Honjo MN, Kudoh H (2016) RNA-Seq reveals virus-virus and virus-plant interactions in nature. FEMS Microbiol Ecol 92(11):pii: fiw176. https://doi.org/10.1093/femsec/fiw176 Melcher U (1989) Symptoms of Cauliflower mosaic virus infection in Arabidopsis thaliana and turnip. Bot Gaz 150:139–147 Narabayashi T, Iwahashi F, Kaido M, Okuno T, Mise K (2009) Melandrium yellow fleck bromovirus infects Arabidopsis thaliana and has genomic RNA sequence characteristics that are unique among bromoviruses. Arch Virol 154(9):1381–1389 Vainio EJ, Chiba S, Ghabrial SA, Maiss E, Roossinck M, Sabanadzovic S, Suzuki N, Xie J, Nibert M, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Partitiviridae. J Gen Virol 99:17–18

A

160

Arabis hirsuta (Hairy rockcress)

Arabis hirsuta (Hairy rockcress) Family: Brassicaceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Weed host

(ArMV)

Family: Secoviridae

Geographical distribution ArMV was first reported infecting Arabis hirsuta in England (Smith and Markham 1944). Symptoms and host(s) The virus-infected hairy rockcress plants exhibit symptoms of mosaics, mottling, chlorotic ringspots, and sometimes necrosis. The virus infects a wide range of weeds and crop plants (Pitcher et al. 1974). Transmission This virus is transmitted by nematode vectors (Xiphinema diversicaudatum, X. bakeri, X. coxi, and Longidorus caespiticola) in a non-persistent manner (Valdy 1972; Brown and Trudgill 1998). The virus is also transmitted by mechanical sap-inoculation, and by grafting (Murant 1983). Virion properties and genome The virions are isometric, non-enveloped of two types but similar in size, 25–30 nm in diameter, and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is bipartite, linear, positive-sense, single-stranded RNA. RNA1 consists of 7334 nt (AY303786 = NC_006057) and RNA2 of 3820 nt (AY017339 = NC_006056) (Loudes et al. 1995; Wetzel et al. 2001, 2004; Dunez and Le Gall 2011; Sanfacon 2015; Thompson et al. 2017).

References Brown DJF, Trudgill DL (1998) Nematode transmission of plant viruses: a 30 year perspective. Host pathogen interactions and crop protection. SCRI Annual Report, pp 121–125 Dunez J, Le Gall O (2011) Nepovirus. Comoviridae. In: The Springer index of viruses. Springer, New York, pp 361–369. https://doi.org/10.1007/978-0-387-95919-1_55 Loudes AM, Ritzenthaler C, Pinck M, Serghini MA, Pinck L (1995) The 119 kDa and 124 kDa polyproteins of Arabis mosaic nepovirus (isolate S) are encoded by two distinct RNA2 species. J Gen Virol 76:899–906 Murant AF (1983) Seed and pollen transmission of nematode-borne viruses. Seed Sci Technol 11:973 Pitcher RS, Siddiqi MR, Brown DJF (1974) Xiphinema diversicaudatum. C.I.H. Descriptions of plant-parasitic nematodes, Set 4, No. 60. 4 pp Sanfacon H (2015) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. In: eLS. Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000764.pub3. http://www.els.net Smith KM, Markham R (1944) Two new viruses affecting tobacco and other plants. Phytopathology 34:324–329 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J Gen Virol 98:529–531 Valdy RB (1972) Transmission of Raspberry ringspot virus by Longidorus caespiticola, L. leptocephalus and Xiphinema diversicaudatum and of Arabis mosaic virus by L. caespiticola and X. diversicaudatum. Ann Appl Biol 71:229–234 Wetzel T, Meunier L, Jaeger U, Reustle GM, Krczal G (2001) Complete nucleotide sequences of the RNAs 2 of German isolates of grapevine fanleaf and Arabis mosaic nepoviruses. Virus Res 75(2):139–145 Wetzel T, Beck A, Wegener U, Krczal G (2004) Complete nucleotide sequence of the RNA 1 of a grapevine isolate of Arabis mosaic virus. Arch Virol 149(5):989–995

Arachis hypogaea (Peanut/groundnut)

161

Arachis glabrata (Rhizoma peanut) Family: Fabaceae

Peanut mottle virus Taxonomic position Genus: Potyvirus

A

Fodder

(PeMoV)

Family: Potyviridae

PeMoV infection in plants of Arachis glabrata was reported from North America (Nischwitz et al. 2007). The virus-infected rhizoma peanut plants exhibit chlorotic ringspot symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PeMoV, refer to Arachis hypogaea.

Peanut stunt virus Taxonomic position Genus: Cucumovirus

(PSV)

Family: Bromoviridae

PSV infection in plants of Arachis glabrata was reported from Florida and Georgia, USA (Blount et al. 2002). The virus-infected rhizoma peanut plants exhibit symptoms of malformed leaves, plant chlorosis, leaf mottling, and stunted plants, which resulted in reduced foliage yield. The virus is transmitted by aphids in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of PSV, refer to Arachis hypogaea.

References Blount AR, Pittman RN, Smith BA, Morgan RN, Dankers W, Sprenkel RK, Momol MT (2002) First report of Peanut stunt virus in perennial peanut in North Florida and Southern Georgia. Plant Dis 86:326 Nischwitz C, Maas AL, Mullis SW, Culbreath AK, Gitaitis RD (2007) First report of Peanut mottle virus in forage peanut (Arachis glabrata) in North America. Plant Dis 91:632

Arachis hypogaea (Peanut/groundnut) Family: Fabaceae

Oil seed

Bean common mosaic virus Synonyms Peanut stripe virus (PStV)

(BCMV)

162

Taxonomic position Genus: Potyvirus

Arachis hypogaea (Peanut/groundnut)

Family: Potyviridae

PStV infection in plants of Arachis hypogaea was first reported from China. BCMV infection in Arachis hypogaea plants was reported from Argentina and Korea (Koo et al. 2002; Choi et al. 2006; de Breuil et al. 2015). This virus is present on groundnut in most areas of Brazil, southern East Asia, China, India, Indonesia, Japan, Malaysia, the Philippines, Myanmar, Thailand, Georgia, Taiwan, Vietnam, Korea, Argentina, and the USA (Demski et al. 1984a, 1988; Kuhn et al. 1984; Reddy et al. 1985d; Wongkaew et al. 1988; Ohki et al. 1989; Prasada Rao et al. 1988, 1989; Saleh et al. 1989; Flasinski et al. 1996; Weibo et al. 1997; McDonald et al. 1998; Higgins et al. 1999; Choi et al. 2001; Sharma et al. 2014; Xu et al. 2014; de Breuil et al. 2015). The virus-infected peanut plants exhibit initial symptoms of distinct discontinuous stripes or blotches along the lateral veins on younger leaflets of peanut. Older leaflets show conspicuous mosaic in the form of green islands or oakleaf patterns, and unlike the symptoms of peanut mottle, these symptoms persist in older leaflets (Wongkaew and Dollet 1990; Sreenivasulu et al. 1992). The necrotic strain reported from Taiwan produces necrotic lesions on leaves. The necrosis leading to stunting, severe mosaic and systemic foliar distortion or stripe symptoms (Chang et al. 1990). The natural hosts of this virus include groundnut, cowpea, soybean, Sesamum, Dolichos lablab, and a few legume weeds, including Centrosema pubescens, C. macrocarpum, Calopogonium caeruleum, Crotalaria striata, Desmodium siliquosum, and Pueraria phaseoloides (Mishra et al.1993). The virus is transmitted by aphid vectors Aphis craccivora, A. gossypii, and Myzus persicae, in a non-persistent manner (Sreenivasulu and Demski 1988). Under experimental conditions the virus is seed-borne in groundnut up to c. 60% (Demski et al. 1984b; Demski and Lovell 1985; Ohki et al. 1989; Matsumoto et al. 1991; Xu et al. 1991; Demski et al. 2004).The causal virus is easily sap-transmissible, and experimentally the virus infects 15 legume spp. and 8 nonlegume spp. Chlorotic lesions are produced on Chenopodium amaranticolor and C. quinoa (Demski et al. 1988, 1993; Gunasinghe et al. 1995; Jain et al. 2000; Xu et al. 2014). For more details of BCMV, refer to Phaseolus vulgaris.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV-infected plants of Arachis hypogaea were reported from the USA (Bays and Demski 1986). The virus-infected groundnut plants exhibit initial symptoms of chlorotic rings and spots on diseased leaves, and after 2–3 weeks, the symptoms disappear. The virus is transmitted by the aphid vector, Aphis craccivora, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Capsicum chlorosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(CaCV)

Family: Tospoviridae

CaCV infection in plants of Arachis hypogaea was reported from China and India (Chen et al. 2007; Vijayalakshmi et al. 2016). The virus-infected groundnut plants exhibit symptoms mimicking those

Arachis hypogaea (Peanut/groundnut)

163

caused by Peanut bud necrosis virus (PBNV). The symptoms included mild chlorosis of leaves, chlorotic and necrotic spots and rings on the leaves, and leaflet necrosis. Occasionally, the infected plants show necrosis of growing tips leading to bud necrosis. New leaves are smaller and internodes are reduced in length. Pods produced on the infected plants show discoloration with concentric rings on the surface (Persley et al. 2006). The virus is transmitted by a thrips vector, Ceratothripoides claratris, in a persistent-propagative manner. The first-instar larvae acquire virus from viruliferous plants, and, after a latent period may be transmitted by second-instar larvae and adults. Adult male and female C. claratris did not differ in transmission competency. The virus is mechanically sap-transmissible and has a wide host range. For more details of CaCV, refer to Capsicum annuum.

Cowpea aphid-borne mosaic virus Taxonomic position Genus: Potyvirus

(CABMV)

Family: Potyviridae

CABMV infection in plants of Arachis hypogaea was reported from Brazil (Pio-Ribeiro et al. 2000; Gillaspie et al. 2001). The virus-infected groundnut plants exhibit symptoms of ring-spots and blotches. The virus is transmitted by aphid vectors, Toxoptera citricidus and Aphis gossypii, in a non-persistent manner, and also by mechanical sap-inoculation. The virus is seed-transmitted in Arachis hypogaea to the extent of 0.15% (Gillaspie et al. 2001). For more details of CABMV, refer to Vigna unguiculata.

Cowpea chlorotic mottle virus Taxonomic position Genus: Bromovirus

(CCMV)

Family: Bromoviridae

CCMV infection in plants of Arachis hypogaea was reported from Georgia, USA (Kuhn and Demski 1987). The virus-infected groundnut plants do not exhibit any symptoms. The virus is transmitted by beetle vectors and also by mechanical sap-inoculation. For more details of CCMV, refer to Vigna unguiculata.

Cowpea mild mottle virus Taxonomic position Genus: Carlavirus

(CPMMV)

Family: Betaflexiviridae

CPMMV infection in plants of Arachis hypogaea was reported from India, Kenya, and Sudan (Iizuka et al. 1984; Iwaki et al. 1986; Mali and Nirmal 1987; Nolt and Rajeshwari 1987; Sivaprasad and Sreenivasulu 1996; El-Hassan et al. 1997; Mukoye et al. 2015). The virus-infected groundnut plants exhibit symptoms of vein-clearing and downward rolling of the leaves followed by necrosis and severe stunting. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of CPMMV, refer to Vigna unguiculata.

A

164

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Arachis hypogaea (Peanut/groundnut)

(CMV)

Family: Bromoviridae

CMV infection in plants of Arachis hypogaea was reported from Korea, China, and Argentina (Xu and Barnett 1984; De Breuil et al. 2005, 2008; Park et al. 2015). The virus-infected groundnut plants exhibit chlorotic spots and upward rolling of young leaflets. Adjacent spots may coalesce to form large yellow blotches. Subsequently developed leaflets exhibit yellowing of the lamina with green stripes along the lateral veins. Occasionally, leaflets are deformed and plants are moderately stunted. The virus is transmitted by more than 75 aphid species in a non-persistent manner. The virus is seed-transmitted in groundnut up to 2–4% (Xu and Barnett 1984). The virus is mechanically sap-transmissible to a number of host plants. For more details of CMV, refer to Cucumis sativus.

Groundnut bud necrosis orthotospovirus

(GBNV)

Synonyms Peanut bud necrosis virus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

Geographical distribution Bud necrosis of Arachis hypogaea plants was first reported from India by Reddy et al. (1968). GBNV occurs in South and Southeast Asia including Bangladesh, China, India, Iran, Vietnam, Nepal, Pakistan, Sri Lanka, Taiwan, Indonesia, Thailand, Ivory Coast, and the Philippines (Reddy et al. 1992, 1995; Delfosse et al. 1995; Wongkaew and Chuapong 1997; Golnaraghi et al. 2002; Damayanti and Naidu 2009; Vemana and Jain 2012; Basavaraj et al. 2017). Symptoms and host(s) The virus-infected groundnut plants exhibit the initial symptoms of chlorotic spots, and necrotic lesions on young quadrifoliates. These lesions cover the entire leaflet, leading to complete necrosis of young quadrifoliates. These symptoms will be followed by necrosis of the entire stem located below the necrosed quadrifoliates. If young plants are affected (less than 1-month-old), the entire plant is often necrosed. In the case of older plants, one or more branches will show necrosis. These plants are stunted and often show axillary shoot proliferation as in the case of peanut bud necrosis disease. Only apical branches show some axillary shoot proliferation. Terminal bud necrosis often occurs when temperatures are relatively high. Necrotic spots are observed on the majority of pods. The size of the pods is severely reduced and kernels are not marketable. The virus infects several crops such as mungbean, urdbean, cowpea, soybean, sunflower, carrot, tomato, potato, cotton, and Lablab purpureus (Jain 2004; Holkar et al. 2017). Transmission The virus is transmitted by thrips vectors, Thrips palmi, Frankliniella schultzei, and Scirtothrips dorsalis, in a persistent-propagative manner (German et al. 1992; Vijayalakshmi 1994; Lakshmi et al. 1995; Daimei et al. 2017). The virus survives in a number of hosts that act as a source of inoculum for the vector. The thrips are carried by wind. The population of vectors increases rapidly from

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January–March and August–September to Autumn, and hence the crop suffers a heavy loss in both the seasons. A prolonged dry spell favors the multiplication of thrips and spread of the virus. The virus is mechanically sap-transmissible to a wide host range, over 600 plant species in more than 70 families. Diagnostic hosts are Vigna unguiculata (concentric chlorotic and necrotic rings being formed on leaves of cvs. C-152 or California black eye 4–5 days after inoculation) and Petunia hybrida (necrotic lesions appearing on leaves within 3–4 days). The virus is not transmitted through peanut seed (Reddy et al. 1995). Virion properties and genome The virions are spherical and membrane bound, of 80–120 nm in diameter. The genome is comprised of three unique molecules of negative or ambisense ssRNA species: small (S) RNA of 3057 nt (U27809 = NC_003619), medium (M) RNA of 4801 nt (U42555 = NC_003620), and large (L) RNA of 8911 nt (AF025538) (Satyanarayana et al. 1996a, b; Gowda et al. 1998).

Groundnut chlorotic fan-spot virus

(GCFSV)

Taxonomic position GCFSV is a tentative member of the genus Orthotospovirus and family Tospoviridae. Geographical distribution GCFSV infection in plants of Arachis hypogaea was reported from Taiwan (Chen and Chiu 1995, 1996; Chu et al. 2001). Symptoms and host(s) The virus on peanut is characterized by large chlorotic, fan-shaped spots with conspicuous concentric rings that often occurred in succession along the main veins of the leaflets. Affected areas of leaves later turned bright yellow, and finally brown and necrotic. Transmission The virus is transmitted by a thrips vector, Scirtothrips dorsalis, in a persistent-propagative manner. The virus is also mechanically sap-transmissible to a number of hosts (Chen and Chiu 1996; Chen et al. 1996). Virion properties and genome The virions are quasi-spherical, enveloped particles with diameters of 80–100 nm and a tripartite singlestranded RNA genome (Chu et al. 2001). Sequences of the L RNA (KP146140; 8796 nt) and M RNA (KP146141; 4857 nt) are available.

Groundnut ringspot orthotospovirus Taxonomic position Genus: Orthotospovirus

(GRSV)

Family: Tospoviridae

A

166

Arachis hypogaea (Peanut/groundnut)

Geographical distribution GRSV infection was first reported in Arachis hypogaea plants from South Africa by Klesser (1966). The virus spreads in Argentina, the Netherlands, Brazil, South America, Florida (USA), and Africa (De Breuil et al. 2007, 2008; McAvoy et al. 2011; Camelo-Garcia et al. 2014). Symptoms and host(s) The virus-infected groundnut plants exhibit symptoms of ring-spots in leaves. Transmission The virus is transmitted by thrips vectors, Frankliniella occidentalis, F. intonsa, F. gemina, and F. schultzei in a persistent and propagative manner (Wijkamp et al. 1995; Nagata et al. 2004). The virus is mechanically sap-transmissible. Virion properties and genome The virions are spherical and membrane bound 80–120 nm in diameter. The genome is comprised of three unique molecules of negative or ambisense single-stranded RNAs (King et al. 2012). Sequences of the L RNA (KY350136; 8876 nt), M RNA (KY350137; 4925 nt) and S RNA (KY400110; 3069 nt) of different isolates are available.

Groundnut rosette assistor virus Taxonomic position Genus: Unassigned

(GRAV)

Family: Luteoviridae

Geographical distribution GRAV infection in plants of Arachis hypogaea was reported from Africa (Hull and Adams 1968; Reddy et al. 1985a; Naidu et al. 1999; Anitha et al. 2014; Okello et al. 2014). Groundnut rosette unbravirus is the most important virus disease of groundnuts in Africa, and requires GRAV for aphid transmission. Found, but with no evidence of spread, in India and the Philippines. Symptoms and host(s) The virus infection alone is asymptomatic in groundnut. The only natural host of this virus is peanut (Arachis hypogaea). Transmission The virus is transmitted by an aphid vector, Aphis craccivora, in a circulative non-propagative manner. The virus is retained when the vector moults, does not multiply in the vector, is not transmitted congenitally to the progeny of the vector, but can help the vector transmission of another virus (Groundnut rosette umbravirus). The virus is not transmissible by mechanical inoculation, by grafting, by contact between plants, by seed, or by pollen. Virion properties and genome The virions are isometric, non-enveloped, 25 nm in diameter, and rounded in profile with a conspicuous capsomere arrangement (Rajeshwari and Murant 1988). The genome consists of a positive-sense single-stranded RNA of which only fragments are available, including ORF 3 (790 nt; KX607035) and coat protein gene of 600 nt (Z68894, AF195824) are available. (AF195824) (Deom et al. 2000).

Arachis hypogaea (Peanut/groundnut)

Groundnut rosette virus Taxonomic position Genus: Umbravirus

167

(GRV)

Family: Tombusviridae

Geographical distribution GRV infection in plants of Arachis hypogaea was first reported in 1907 from Tanzania (Zimmerman 1907). Since then this virus has been reported in sub-Saharan Africa and several African countries that include Malawi, Nigeria, Uganda, Senegal, Burkina Faso, Cote d’Ivoire, South Africa, Niger, and Kenya (Bisht et al. 1963; Dollet et al. 1986; Naidu et al. 1999; Taliansky et al. 2000; Wangai et al. 2001; Anitha et al. 2014; Okello et al. 2014; Mugisa et al. 2016).

Symptoms and host(s) The virus-infected peanut plants show symptoms in three predominant forms, viz., chlorotic rosette, green rosette, and mosaic rosette. The chlorotic rosette is ubiquitous in sub-Saharan Africa, while green rosette is distributed in West Africa, Uganda, and Angola, and mosaic rosette is recorded only in East and Central Africa (Storey and Ryland 1957; Reddy et al. 1985b). These symptoms differ with the strains, but stunting and rosette are always present. The number of seeds and pods are severely decreased (20–80%). The only known natural host of this virus is peanut (Arachis hypogaea) (Okusanya and Watson 1966).

Transmission The virus is transmitted by an aphid vector Aphis craccivora in a circulative, non-propagative manner (Storey and Ryland 1955; Hull 1964; Misari et al. 1988) in the presence of Groundnut rosette assistor virus. The virus is retained when the vector moults. It does not multiply in the vector, and is not transmitted congenitally to the progeny of the vector. Groundnut rosette disease is caused by a complex of three agents: Groundnut rosette assistor virus (GRAV) (Hull and Adams 1968; Scott et al. 1996), Groundnut rosette virus (GRV) (Murant 1990; Taliansky et al. 1996), and the satellite RNA (sat-RNA) (Murant et al. 1988) of GRV. The Groundnut rosette virus (GRV) depends on Groundnut rosette assistor virus (GRAV) for transmission by Aphis craccivora. The main cause of rosette disease symptoms in plants is due to a third component, a satellite RNA, which depends on GRV for its replication but on GRAV for its encapsidation and transmission by aphids. The aphid may transmit GRAVonly, GRVand satellite RNA, but not GRV alone (Deom et al. 2000). GRV is mechanically sap-transmissible but not transmissible by contact between plants. No seed transmission is recorded.

Virion properties and genome No virus-like particles have been seen in preparations from plants infected only with GRV, and no candidate for the role of a particle protein has been identified among the proteins putatively encoded by the genomic RNA. The virus seems to exist in plants as an infective ssRNA. The genome is singlestranded, positive-sense RNA, of 4019 nt (Z69910 = NC_003603) (Casper et al. 1983; Breyel et al. 1985; Reddy et al. 1985b; Taliansky et al. 1996).

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Arachis hypogaea (Peanut/groundnut)

Groundnut yellow spot orthotospovirus

(GYSV)

Synonyms Peanut yellow spot virus (PYSV) Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

Geographical distribution GYSV infection in plants of Arachis hypogaea was first reported from India in 1979 (Anon 1980). The virus was also reported from Thailand and Myanmar (Wongkaew 1986; Reddy et al. 1991; Satyanarayana et al. 1996c; Basavaraj et al. 2017). Symptoms and host(s) The virus in groundnut is characterized by yellow spots followed by necrosis of leaves. The virus symptoms begin with chlorotic/yellow leaf spots, which later coalesce and become necrotic (Satyanarayana et al. 1996c). Transmission The virus is transmitted by a thrips vector, Scirtothrips dorsalis, in a persistent-propagative manner (Reddy et al. 1991). Larvae could acquire the virus in 30 min, and the maximum percentage transmission of 43.8% by individual insects resulted following 2-day AAP. A single adult thrip transmitted the virus after a minimum IAP of 30 min. The percentage transmission (33.3%) increased linearly with an increase in IAP up to 1.5 days and a maximum of up to 55 h of IAP (36.1%). GYSV was persistently transmitted for more than 75% of their life-span (Gopal et al. 2010). The virus is sap-transmissible and became systemic in pea, mungbean, and cowpea at temperatures between 20  C and 30  C (Reddy et al. 1991). Virion properties and genome The virions are spherical and membrane bound 80–120 nm in diameter. The coat protein has a molecular mass of 29 kDa. The genome comprises three unique molecules of negative or ambisense ssRNA, a large (L) RNA (c.8900 nt), a medium (M) RNA (c.4800 nt), and a small (S) RNA (c.3000 nt) (AF013994; HQ402596). In addition, a fourth RNA species of approximately 1800 nucleotides was also present in purified preparations (Satyanarayana et al. 1996c, 1998).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Arachis hypogaea was reported from the USA (Pappu et al. 1999; Wells et al. 2001). The virus-infected groundnut plants exhibit symptoms of yellowing, wilting, and necrosis of taproot and crown. The virus is transmitted by a thrips vector Frankliniella occidentalis in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Arachis hypogaea (Peanut/groundnut)

Indian peanut clump virus Taxonomic position Genus: Pecluvirus

169

(IPCV)

Family: Virgaviridae

Geographical distribution Clump disease was first reported in 1927 in a groundnut crop in India (Sundararaman 1927). IPCV infection in plants of Arachis hypogaea was reported from India and Pakistan (Reddy et al. 1983, 1999). Symptoms and host(s) The virus-infected young leaflets of groundnut show mottling, mosaic, and chlorotic rings. When leaflets mature, they turn green, with or without faint mottling. Plants infected at early growth stages become severely stunted (Nolt et al. 1988). The virus infects a number of cereal crops and graminaceous weeds, some symptomlessly and others in which infection induces stunting (Doucet et al. 1999). Transmission The virus is transmitted by a fungal vector Polymyxa graminis (Ratna et al. 1991; Delfosse et al. 2002). The virus is mechanically sap-transmissible and has a wide host range which includes both monocotyledons and dicotyledons (Reddy et al. 2005). From India, Reddy et al. (1988, 1998) have reported seed transmission between 3.5% and 17% in peanut with Indian strain. Virion properties and genome The virions are rod-shaped, of about 24 nm in diameter and of two predominant lengths of 180 and 250 nm. Virions have helical symmetry with a pitch of 2.6 nm. The virus contains a single capsid protein of 24 kDa. The genome consists of two molecules of linear positive single-stranded RNA. RNA1 is 5841 nt (X99149 = NC_004729) contains two ORFs and RNA2 of 4507 nt (AF447397 = NC_004730) contains five ORFs (Reddy et al. 1985c; Miller et al. 1996; Naidu et al. 1996, 2003; Erhardt and Bouzoubaa 2011; Adams et al. 2017).

Mungbean yellow mosaic virus Taxonomic position Genus: Begomovirus

(MYMV)

Family: Geminiviridae

MYMV infection in plants of Arachis hypogaea was reported from India (Sudhakar Rao et al. 1980). The virus-infected groundnut plants exhibit symptoms of bright yellow patches on the leaves with shortening, thickening, and slight upward curling of leaves. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmitted by mechanical sap-inoculation. For more details of MYMV, refer to Vigna radiata.

Passion fruit woodiness virus Taxonomic position Genus: Potyvirus

(PWV)

Family: Potyviridae

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Arachis hypogaea (Peanut/groundnut)

PWV infection in plants of Arachis hypogaea was reported from Australia (Boswell and Gibbs 1983). The virus-infected groundnut plants display dead and yellow areas or streaks, mottling, and wrinkling. The virus is transmitted by aphid vectors Aphis gossypii and Myzus persicae in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PWV, refer to Passiflora edulis.

Peanut chlorotic streak virus Taxonomic position Genus: Soymovirus

(PCSV)

Family: Caulimoviridae

Geographical distribution PCSV infection in plants of Arachis hypogaea is widely distributed in India (Reddy et al. 1993; Satyanarayana et al. 1994a). Symptoms and host(s) The virus-infected peanut plant leaves have oval, chlorotic streaks along the veins on the younger leaflets. Streaks are not distinct in older leaflets and are seen only when viewed against light; infected plants are stunted. The natural host range of this virus is restricted largely to Leguminosae and Solanaceae. Transmission There is no known vector for this virus. The virus is transmitted by mechanical sap-inoculation, and also by grafting. The virus is not transmitted by seed. Virion properties and genome The virions are isometric, non-enveloped, and 42–50 nm in diameter with no obvious surface structure. The genome consists of a single molecule of non-covalently closed circular dsDNA of 8174 bp (U13988 = NC_001634) (Richins et al. 1990; Satyanarayana et al. 1994b; Hibi 2011).

Peanut clump virus Taxonomic position Genus: Pecluvirus

(PCV)

Family: Virgaviridae

Geographical distribution PCV infection in plants of Arachis hypogaea was reported from West Africa (Thouvenel et al. 1976, 1988; Dollet et al. 1986; Delfosse et al. 1995). Symptoms and host(s) Peanut clump disease is characterized by patches of stunted or clumped groundnut plants. The virusinfected groundnut plants usually have upright stems and short internodes with leaflets that are smaller and darker green in color. A yellowing strain causes patterns of concentric yellow lines in older leaves, but no stunting (Manohar et al. 1995).

Arachis hypogaea (Peanut/groundnut)

171

Transmission The virus is transmitted by a soil-borne fungal vector, Polymyxa graminis (Dieryck et al. 2008, 2011). The virus is transmitted by mechanical sap-inoculation to several dicots and monocots including species in the Aizoaceae, Amaranthaceae, Chenopodiaceae, Cucurbitaceae, Gramineae, Leguminosae, Scrophulariaceae, and Solanaceae. The virus is also transmitted by seed at levels of 3.5–17% in Arachis hypogaea (Thouvenel and Fauquet 1981). Virion properties and genome The virions are rod-shaped of about 21 nm in diameter and of two predominant lengths of 190 and 245 nm. Virions have helical symmetry with a pitch of 2.6 nm. The genome consists of two molecules of linear positive-sense ssRNA: RNA1 is 5897 nt (X78602 = NC_003672) contains two ORFs and RNA2 of 4504 nt (L07269 = NC_003668) contains five ORFs (Manohar et al. 1993; Herzog et al. 1994; Naidu et al. 2003; Erhardt and Bouzoubaa 2011; Adams et al. 2017).

Peanut mottle virus

(PeMoV)

Synonyms Peanut mild mosaic virus; Peanut severe mosaic virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution PeMoV infection in plants of Arachis hypogaea was first reported from the USA in 1961 (Bock and Kuhn 1975; Kuhn and Demski 1984). The virus is widely distributed in groundnut-growing countries (Bock 1973; Paguio and Kuhn 1973; Reddy et al. 1978, 1985d; Behncken 1980; Ahmed and Idris 1981; Kuhn et al. 1984; Porter et al. 1984; Highland et al. 1988; Sreenivasulu et al. 1988; Koo et al. 2002; Spiegel et al. 2008; Beikzadeh et al. 2015). Symptoms and host(s) The virus-infected groundnut plants exhibit symptoms varying from mild mottle to definite mottle or mosaic, sometimes with necrosis. The range of symptoms includes mild mottle on youngest leaflets; upward curling of edges of older leaflets show interveinal depression, and mild mottling. Some genotypes may not show upward curling of leaf edges. In some genotypes interveinal depressions and inward rolling of margins of leaflets can occur. Differences in symptoms are due to the presence of different virus strains and probably different host cultivars and environmental conditions (Reddy and Demski 1996). The virus has a moderate host range; Pisum sativum is used as a virus propagation host and Phaseolus vulgaris cv. Topcrop as an assay host. Transmission The virus is transmitted by aphid vectors, viz., Aphis craccivora, A. gossypii, Hyperomyzus lactucae, Myzus persicae, Rhopalosiphum padi, and R. maidis in a non-persistent manner (Demski 1975; Highland et al. 1981; Sreenivasulu and Demski 1988; Reddy and Demski 1996). The virus is transmitted by mechanical sap-inoculation to 27 legumes (among which are several important legume crops), and 4 non-legumes have been reported as experimental hosts. Seed transmission frequency is 1–8.5% in peanut (Adams and Kuhn 1977; Reddy et al. 1984; Puttaraju et al. 2001).

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Virion properties and genome The virions are non-enveloped, flexuous filaments, 750 nm long and 12 nm wide. The genome consists of a single molecule of linear, positive-sense, single-stranded RNA of 9709 nt (AF023848 = NC_002600) (Sun and Hebert 1972; Rajeshwari et al. 1983; Tolin and Ford 1983; Revers and Garcia 2015; Wylie et al. 2017).

Peanut stunt virus Taxonomic position Genus: Cucumovirus

(PSV)

Family: Bromoviridae

Geographical distribution PSV infection in plants of Arachis hypogaea was reported from Europe, Japan, Korea DPR (North), Korea Republic, Morocco, Poland, Spain, Sudan, China, Georgia, Iran, and the USA (Miller and Troutman 1966; Mink et al. 1969; Kuhn et al. 1984; Tolin 1984; Ahmed 1985; Zeyong et al. 1998; Amid-Motlagh et al. 2017). Symptoms and host(s) The virus-infected groundnut plants exhibit symptoms of paler green and/or yellowed leaves, which are malformed. The pods of plants infected with the virus are frequently small and malformed, and the shells are commonly split open to expose seed (Mink et al. 1969). Transmission The virus is transmitted by aphid vectors, Aphis craccivora, A. spiraecola, and Myzus persicae in a nonpersistent manner. The virus is transmitted by mechanical sap-inoculation, and is experimentally infectious to a wide range of non-leguminous plants (Fischer and Lockhart 1978). Seed transmission of this virus in groundnut ranges between 3% and 4% (Troutman et al. 1967). Virion properties and genome The virions are non-enveloped, spherical of about 25–30 nm in diameter with T = 3 icosahedral symmetry, and composed of 180 coat proteins: 12 pentamers and 20 hexamers. The genome is segmented and tripartite linear positive-sense ssRNA, composed of RNA1 which is 3357 nt (U15728 = NC_002038), RNA2 of 2947 nt (U15729 = NC_002039), and RNA3 of 2188 nt (U15730 = NC_002040). Each genomic segment has a 30 tRNA-like structure and a 50 cap (Mink et al. 1969; Karasawa et al. 1991, 1992; Naidu et al. 1995; Zeyong et al. 1998; Garcia-Arenal Rodriguez and Fraile 2011; Scott 2011).

Peanut yellow mosaic virus

(PeYMV)

Synonyms Peanut yellow mottle virus Taxonomic position Genus: Tymovirus

Family: Tymoviridae

Arachis hypogaea (Peanut/groundnut)

173

Geographical distribution PeYMV infection in plants of Arachis hypogaea was reported from India and Nigeria (Lana 1980; Reddy and Demski 1996; Kumar et al. 2007).

Symptoms and host(s) The virus-infected groundnut plants exhibit mosaic mottle and yellowing symptoms.

Transmission The virus is transmitted by beetle vectors in a semi-persistent manner. The virus is also mechanically sap-transmissible to species belonging to the families Fabaceae, Chenopodiaceae, and Solanaceae.

Virion properties and genome The virions are non-enveloped, isometric capsid with T = 3 icosahedral symmetry, about 30 nm in diameter. The genome consists of a single molecule of positive-sense ssRNA. The 30 -terminus has a tRNA-like structure (Lana 1980; Martelli et al. 2002). No sequence is currently available.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Arachis hypogaea was reported from India, Pakistan, South Africa, North American region and the Pacific region, Australia, Brazil, Canada, Peru, and the USA (Costa and Carvalho 1961; Reddy and Demski 1996; Cook et al.1999; Reddy et al. 2001, 2002; Prasada Rao et al. 2005; Vemana and Jain 2012; Sharman et al. 2015). The virus-infected groundnut plants exhibit symptoms initially on young leaves, as necrotic lesions and veinal which later spreads to the petiole and stem. Necrotic lesions on the stem later spread upward, killing the bud. In some cultivars of axillary shoots is noticed. In the late infected plants, only terminal leaflets or a single branch may show necrosis. In some cultivars necrotic spots appear on the pegs and pods. In severe infection the entire plant becomes necrotic and dies prematurely. The virus is transmitted by thrips vectors, Frankliniella occidentalis, F. schulzei, Thrips tabaci, Scirtothrips dorsalis, and Megalurothrips usitatus, and among these thrips vectors, F. schulzei is more efficient than other species in disease transmission. The virus present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is transmitted by mechanical sap-inoculation, and by grafting. The seed infection ranged from 18.9% to 28.9% among the seeds collected from naturally infected and sap-inoculated groundnut varieties (JL 24, TMV 2, Prasuna, Kadiri 6, Kadiri 9, Anantha, and Kadiri 7 Bold) belonging to Spanish and Virginia types. Further, TSV was detected both in pod shell and seed testa, and none of the samples show the presence of TSVeither in cotyledon or embryo. Growout and bioassay tests proved the absence of seed transmission in groundnut and other legume crops (Reddy et al. 2002, 2007; Vemana and Jain 2010). For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

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Tomato chlorotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(TCSV)

Family: Tospoviridae

TCSV infection in plants of Arachis hypogaea was reported from Haiti (Adegbola et al. 2016). The virus-infected groundnut plants exhibit symptoms of chlorosis, necrosis, distortion and mottling of young leaves, necrosis of the terminal buds, axillary shoot proliferation, and stunting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also mechanically saptransmissible. For more details of TCSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWVoccurrence on Arachis hypogaea plants was first reported in Brazil (Costa 1941). The virus also occurs in North America, South America, Sri Lanka, Iran, and Africa (Dollet et al. 1986; Jayasena et al. 1988; Sreenivasulu et al. 1988; Hagan et al. 1990; Porter et al. 1991; Sreenivasulu et al. 1991; Jain et al. 1997, 1998; Golnaraghi et al. 2001; Culbreath et al. 2003; Culbreath and Srinivasan 2011). The virus causes a wide range of symptoms on peanut, from local lesions (chlorotic and/or necrotic) to systemic symptoms such as mild mottling and bronzing, to striking geometric rings on foliage, to severe stunting of the entire plant, to necrotic streaks on the stem, and, in severe cases, to plant wilt and death. Peanut pegs, pods, and kernels also may be stunted, deformed, and discolored. The virus is transmitted by thrips vector, viz., Thrips tabaci, T. setosus, T. parmi, Frankliniella schultzei, F. occidentalis, F. fusca, and Scirtothrips dorsalis in a persistent-propagative manner (Palmer et al. 1990; Brown et al. 1996; Garcia et al. 2000). The virus is transmissible by mechanical sapinoculation to a number of herbaceous hosts, transmitted by grafting, and not transmissible by contact between plants. The virus is not transmitted by seed and pollen (Mandal et al. 2001). For more details of TSWV, refer to Solanum lycopersicum.

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Arachis pintoi (Pinto peanut)

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Tolin S (1984) Peanut stunt virus. In: Porter DN, Rodriguez-Kabana R (eds) Compendium of peanut diseases. American Phytopathological Society, St. Paul, pp 45–48 Tolin SA, Ford RH (1983) Purification and serology of Peanut mottle virus. Phytopathology 73:899–903 Troutman JL, Bailey WK, Thomas CA (1967) Seed transmission of Peanut stunt virus. Phytopathology 57:1280–1281 Vemana K, Jain RK (2010) New experimental hosts of Tobacco streak virus and absence of true seed transmission in leguminous hosts. Indian J Virol 21:117–127 Vemana K, Jain RK (2012) Biological similarities and differences between Tobacco streak virus and Groundnut bud necrosis virus infecting groundnut (Arachis hypogea). Indian Phytopathol 65:177–183 Vijayalakshmi K (1994) Transmission and ecology of Thrips palmi Karny, the vector of Peanut bud necrosis virus. Dissertation, Andhra Pradesh Agricultural University Vijayalakshmi G, Haokip BD, Karthikeyan G, Alice D, Ganapathy N, Asokan G, Rajendran L, Kennedy S (2016) First report of field infection of Capsicum chlorosis virus on groundnut (Arachis hypogaea) in India (Tamil Nadu). Plant Dis 100:2339 Wangai AW, Pappu SS, Pappu HR, Deom CM, Naidu RA (2001) Distribution and characteristics of groudnut rosette disease in Kenya. Plant Dis 85:470–474 Weibo D, Dunyu Y, Xingqi G, Xiaoping Z (1997) Effect of PStV infection on physiological and biochemical properties of peanut. Acta Phytopathologica Sinica 27:281–285 Wells ML, Pappu HR, Culbreath AK, Todd JW, Brown SL (2001) Field survey of impatiens necrotic spot virus in Georgia peanut. Peanut Sci 28:34–37 Wijkamp I, Almarza N, Goldbach R, Peters D (1995) Distinct levels of specificity in thrips transmission of tospoviruses. Phytopathology 85:1069–1074 Wongkaew S (1986) Peanut stripe and other viruses in Thailand. In: Proc. Peanut CRSP Workshop. Khon Kaen, Thailand, pp 86–90 Wongkaew S, Chuapong J (1997) Detail study of Groundnut bud necrosis virus isolate. Proceedings of the 12th Thailand National Peanut Meeting, Khon Kaen, Thailand, 25–27 October 1994, Khon Kaen FIELD Crops Research Center, Thailand, pp 214–220 Wongkaew S, Dollet M (1990) Comparison of Peanut stripe virus isolates using symptomatology on particular hosts and serology. Oleagineux 45:267–278 Wongkaew S, Kantrong S, Choopanya D (1988) Peanut stripe virus disease in Thailand. Coordination of research on Peanut stripe virus: summary proceedings of the First Meeting to Coordinate Research on Peanut Stripe Virus Disease of Groundnut, 9–12 June 1987, Malang. ICRISAT, Patancheru, A.P. 502 324, India Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Xu Z, Barnett OW (1984) Identification of a Cucumber mosaic virus strain from naturally infected peanut in China. Plant Dis 68:386–389 Xu Z, Chen K, Zhang Z, Chen J (1991) Seed transmission of Peanut stripe virus in peanut. Plant Dis 75:723–726 Xu M, Gao F, Yang J, Wu J, Xie L, Chi Y (2014) Complete genome sequence of Peanut stripe virus isolated in China. J Phytopathol 162:829–832 Zeyong X, Higgins CM, Kunrong C, Dietzgen RG, Zhongyi Z, Liying Y, Xiaoping F (1998) Evidence for a third taxonomic subgroup of Peanut stunt virus from China. Plant Dis 82:992–998 Zimmerman A (1907) Uber eine krankheit der erdnusse. Translated about a disease (illness) of the groundnut (Arachis hypogaea L.). Pflanzer 3:129–133

Arachis pintoi (Pinto peanut) Family: Fabaceae

Arachis pintoi virus Taxonomic position Genus: Allexivirus

Fodder plant

(ApV)

Family: Alphaflexiviridae

A

182

Aralia spp.

Geographical distribution ApV infection in plants of Arachis pintoi was reported from Colombia (Gutierrez Sanchez et al. 2016). Transmission The virus is transmitted by mechanical sap inoculation. Use of infected vegetative propagative material is another source of virus spread. Virion properties and genome The virions are very flexuous filaments of about 800 nm in length and 12 nm in diameter. The genome contains a single molecule of single-stranded RNA of 7599 nt (KX058345 = NC_032104) and comprises six ORFs, encoding, in order, the replication-related proteins, the first two proteins of a TGB, a serine-rich protein of unknown function, the CP and a putative nucleic acid-binding regulatory protein. Coat protein subunit is of one type, and 28 kDa in size (Gutierrez Sanchez et al. 2016).

Peanut mottle virus Taxonomic position Genus: Potyvirus

(PeMoV)

Family: Potyviridae

PeMoV infection in plants of Arachis pintoi was reported from Brazil and Colombia (Morales et al. 1991; Anjos et al. 1998). The virus-infected pinto peanut plants exhibit foliar chlorotic ring-spot symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PeMoV, refer to Arachis hypogaea.

References Anjos JTN, Kitajima EW, Charchar MJA, Marinho VLA (1998) Natural infection of Arachis pintoi by Peanut mottle virus in central Brazil. Fitopatol Bras 23(1):71–74 Gutierrez-Sanchez PA, Jaramillo-Mesa H, Marin-Montoya MA (2016) Next generation sequence analysis of the forage peanut (Arachis pintoi) virome. Rev Fac Nac Agron Medellin 69:7881–7891 Morales FJ, Castano M, Velasco AC, Arroyave J, Zettler FW (1991) Natural infection of tropical forage legume species of Arachis and Stylosanthes by potyviruses related to Peanut mottle virus. Plant Disease 75:1090–1093

Aralia spp. Family: Araliaceae

Ornamental

Schefflera ringspot virus Taxonomic position Genus: Badnavirus

(SRV)

Family: Caulimoviridae

SRV infection in plants of Aralia spp. was reported from Australia, Barbados, Cuba, Mauritius, Honduras, Taiwan, Thailand, and the USA (Lockhart and Olszewski 1996). The virus-infected aralia

Araujia spp. (Moth plant)

183

plants exhibit symptoms including chlorotic spotting, vein-clearing, and reduction of leaf size. Symptoms occurred sporadically and appeared to be correlated with changes in environmental conditions. The virus is transmitted by the citrus mealybug (Planococcus citri) in a semi-persistent manner. The virus is not transmitted by mechanical inoculation. For more details of SRV, refer to Schefflera arboricola.

References Lockhart BEL, Olszewski NE (1996) Schefflera ring spot virus, the widely distributed mealybug transmitted virus occurring in schefflera, aralia. Acta Hortic 432:196–202

Araujia spp. (Moth plant) Family: Apocynaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Ornamental

(AMV)

Family: Bromoviridae

AMV infection in plants of Araujia sericifera was reported from Italy (Parrella et al. 2013). The virusinfected moth plants exhibit symptoms of bright yellow mosaic or calico stripe and leaf distortion. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a number of test plants. For more details of AMV, refer to Medicago sativa.

Araujia mosaic virus Taxonomic position Genus: Potyvirus

(ArjMV)

Family: Potyviridae

Geographical distribution ArjMV infection in plants of Araujia spp. was reported from Argentina (Charudattan et al. 1980). This virus spreads in the South and Central American region, Argentina, and New Zealand (Hiebert and Charudattan 1984; Elliott et al. 2009). Symptoms and host(s) The virus-infected Araujia angustifolia plants exhibit mosaic symptoms. Transmission The virus is transmitted by aphid vectors, Aphis nerii, Aphis spiraecola, and Myzus persicae, in a nonpersistent manner. The virus is also transmissible by mechanical sap-inoculation to plants of six genera in the Apocynaceae. The virus is not transmissible by contact between plants.

A

184

Arctium lappa (Greater burdock)

Virion properties and genome The virions are non-enveloped, flexuous filaments with a clear modal length of 741 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense, single-stranded RNA. A partial genome sequence is available (EF710625) (Massey et al. 2007; Revers and Garcia 2015; Wylie et al. 2017).

References Charudattan R, Zettler FW, Cordo HA, Christie RG (1980) Partial characterization of a Potyvirus infecting the milkweed vine, Morrenia odorata. Phytopathology 70:909–913 Elliott MS, Massey B, Cui X, Hiebert E, Charudattan R, Waipara N (2009) Supplemental host range of Araujia mosaic virus, a potential biological control agent of moth plant in New Zealand. Australas Plant Pathol 38:603–607 Hiebert E, Charudattan R (1984) Characterization of Araujia mosaic virus by in vitro translation analyses. Phytopathology 74:642–646 Massey B, Cui X, Hiebert E, Elliott MS, Waipara N, Hayes L, Charudattan R (2007) Partial sequencing of the genomic RNA of Araujia mosaic virus and the comparison of the coat protein sequence with other potyviruses. Arch Virol 152:2125–2129 Parrella G, Greco B, Cennamo G, Griffo R, Stinca A (2013) Araujia sericifera new host of Alfalfa mosaic virus in Italy. Plant Dis 97:1387 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Arctium lappa (Greater burdock) Family: Asteraceae

Burdock mottle virus Taxonomic position Genus: Benyvirus

Medicinal

(BdMV)

Family: Benyviridae

Geographical distribution BdMV infection in plants of Arctium lappa was first reported from Japan by Inouye (1973). The virus spreads in Japan (Hirano et al. 1999; Kondo et al. 2013). Symptoms and host(s) The virus-infected Arctium lappa plants exhibit faint leaf chlorosis symptoms. Transmission The virus is transmitted by mechanical sap-inoculation to species in fewer than three families. The mode of transmission in the field is not known. Virion properties and genome The virions are rod-shaped, usually straight, with a clear modal length of 250 nm and 17 nm wide. The genome consists of two segments of positive-sense single-stranded RNA (Hirano et al. 1999; Kondo

Arctium lappa (Greater burdock)

185

et al. 2013). The RNA-1 is 7038 nt (AB818898 = NC_021735) encoding a single protein of 249 kDa. The RNA-2 is 4315 nt (AB818899 = NC_021736) and codes for six proteins (Gilmer et al. 2011, 2017).

Burdock stunt viroid

(BuSVd)

Taxonomic position BuSVD is unassigned viroid which has not been approved as species. BuSVd infection in plants of Arctium lappa was reported from China (Chen et al. 1983). The viroidinfected greater burdock plants exhibit stunting and leaf mottling symptoms. The viroid is mechanically sap-transmissible. For more details of BuSVd, refer to Arctium tomentosum.

Burdock yellows virus Taxonomic position Genus: Closterovirus

(BuYV)

Family: Closteroviridae

Geographical distribution BuYV infection was first reported in Arctium lappa plants from Japan by Inouye and Mitsuhata (1971). The virus spreads in Japan (Nakano and Inouye 1980). Symptoms and host(s) The virus-infected greater burdock plants exhibit symptoms of faint chlorosis, often marginal or symptomless. Transmission The virus is transmitted by an aphid vector, Dactynotus gobonis, in a semi-persistent manner. The virus is also transmitted by mechanical sap-inoculation (with difficulty). Virion properties and genome The virions are flexuous filaments, non-enveloped, with a clear modal length of 1725 nm and 12 nm wide (Nakano and Inouye 1980). The genome is a monopartite, linear, positive-sense, single-stranded RNA of 15,000–20,000 nt (Agranovsky and Lesemann 2011).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Arctium lappa was reported from Iran (Polak and Brcak 1961; Koolivand et al. 2012). The virus-infected greater burdock plants exhibit mild mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

A

186

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

Arctium tomentosum (Woolly burdock)

(TRSV)

Family: Secoviridae

TRSV infection in plants of Arctium lappa was reported from New York (Begtrup 1983; Shiel and Castello 1985). The virus-infected greater burdock plants exhibit ringspot symptoms. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

References Agranovsky AA, Lesemann D-E (2011) Closterovirus. Closteroviridae. In: The Springer index of Viruses. Springer, New York, pp 327–333. https://doi.org/10.1007/978-0-387-95919-1_50 Begtrup J (1983) Arctium lappa L. a new host for Tobacco ringspot virus (TobRV) identified with immunosorbent electron microscopy (ISEM). Tidsskr Planteavl 87:179–182 Chen W, Tien P, Zhu YX, Liu Y (1983) Viroid-like RNAs associated with Burdock stunt disease. J Gen Virol 64:409–414 Gilmer D, Lauber E, Guilley H (2011) Benyvirus. In: Tidona C, Darai G (eds) The Springer index of viruses. Springer, New York. https://doi.org/10.1007/978-0-387-95919-1 Gilmer D, Ratti C, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Benyviridae. J Gen Virol 98:1571–1572 Hirano S, Kondo H, Maeda T, Tamada T (1999) Burdock mottle virus has a high genome similarity to Beet necrotic yellow vein virus. In: Sherwood JL, Rush CM (eds) Proceedings of the fourth symposium of the international working group on plant viruses with fungal vectors, American Society of Sugar Beet Technologists, Denver, pp 33–36 Inouye T (1973) Host range and electron microscopy of Burdock mottle virus, a rod-shaped virus from Arctium lappa L.: studies on the viruses of plants in Compositae in Japan. Ber Ohara Inst Landw Biol, Okayama University 15:207–218 Inouye T, Mitsuhata K (1971) Viruses in burdock, Arctium lappa L. 2. Studies on the viruses of plants in compositae in Japan. Nogaku Kenkyu 54(1):1–14 Kondo H, Hirano S, Chiba S, Andika IB, Hirai M, Maeda T, Tamada T (2013) Characterization of Burdock mottle virus, a novel member of the genus Benyvirus, and the identification of benyvirus-related sequences in the plant and insect genomes. Virus Res 177:75–86 Koolivand D, Bashir NS, Mozafari J (2012) Serological and molecular detection of newly isolated Cucumber mosaic virus variants from Iran. Int J Agric: Res Rev 2:933–941 Nakano M, Inouye T (1980) Burdock yellows virus, a closterovirus from Arctium lappa L. Ann Phytopathol Soc Jpn 46:7–14 Polak Z, Brcak J (1961) Identification of the mosaic of Arctium lappa L. caused by the common Cucumber mosaic virus. Preslia 33:354–358 Shiel PJ, Castello JD (1985) Detection of tobacco mosaic and tobacco ring spot viruses in herbaceous and woody plants near virus-infected white ash trees in central New York. Plant Dis 69:791–795

Arctium tomentosum (Woolly burdock) Family: Asteraceae

Alstroemeria virus X Taxonomic position Genus: Potexvirus

Ornamental

(AlsVX)

Family: Alphaflexiviridae

Arctium tomentosum (Woolly burdock)

187

AlsVX infection in plants of Arctium tomentosum was reported from Finland (Bi et al. 2012). The virusinfected wooly burdock plants show yellow vein symptoms. The virus is mechanically saptransmissible. No vector is known for this virus. For more details of AlsVX, refer to Alstroemeria spp.

Burdock stunt viroid

(BuSVd)

Taxonomic position BuSVD is unassigned viroid which has not been approved as species. Geographical distribution BuSVd infection in plants of Arctium tomentosum was reported from China (Chen et al. 1983, 1986; Chen and Tien 1985). Symptoms and host(s) The viroid-infected wooly burdock plants exhibit stunting and leaf mottling symptoms. Transmission The viroid is mechanically sap-transmissible. Etiology and genome properties Neither virus-like particles nor mycoplasma-like organisms have been detected in the extracts and thin sections of diseased leaves, but two disease-specific RNA species, RNA-1 and RNA-2, were found to be associated with the disease. They had a low molecular weight (RNA-1, 1.8  l05 to 1.9  105; RNA-2, 1.7  l05). Preparations of RNA-1 and RNA-2 contained circular molecular structures.

Woolly burdock yellow vein virus

(WBYVV)

Taxonomic position WBYVV is a tentative member of the genus Emaravirus and family Fimoviridae. Geographical distribution WBYVV infection in plants of Arctium tomentosum was reported from Finland (Bi et al. 2012). Symptoms and host(s) The virus-infected wooly burdock plants show yellow vein symptoms. Transmission The virus is transmitted by mite vectors. Virion properties and genome A partial genome sequence of (“Fig mosaic-like virus JPTV-2012”) WBYVV is available (JQ354894).

A

188

Arctotis spp. (African daisy)

References Bi YQ, Tugume AK, Valkonen JPT (2012) Small-RNA deep sequencing reveals Arctium tomentosum as a natural host of Alstroemeria virus X and a new putative Emaravirus. PLoS One 7(8):e42758 Chen W, Tien P (1985) Study on Burdock stunt viroid (BSV). III. Ultrastructure of diseased leaf tissues. Acta Phytopathol Sin 15:215 Chen W, Tien P, Zhu YX, Liu Y (1983) Viroid-like RNAs associated with burdock stunt disease. J Gen Virol 64:409–414 Chen W, Tien P, Yong XC, Zhu YX, Sun GD (1986) Study of Burdock stunt viroid (BSV). II. The denaturing behavior and occurrence in separate plants of BSV RNA-1 and RNA-2. Sci Sin (Ser B) 26:147

Arctotis spp. (African daisy) Family: Asteraceae

Ornamental

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Arctotis x hybrida was reported from Italy (Parrella et al. 2013). The virusinfected African daisy plants exhibit rapid dieback, generalized reddening, followed by irreversible wilting symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Parrella G, Greco B, Cavicchi L, Bellardi MG (2013) Dieback and wilting caused by Tomato spotted wilt virus in Arctotis  hybrida in Italy. Plant Disease 97:1387–1388

Areca catechu (Areca palm) Family: Arecaceae

Commercial crop

Areca palm velarivirus 1 Taxonomic position Genus: Velarivirus

(ArPV1)

Family: Closteroviridae

Geographical distribution ArPV1 infection in plants of Areca catechu was reported from China (Yu et al. 2015).

Argyranthemum frutescens (Marguerite daisy)

189

Symptoms and host(s) The virus-infected areca palm plants initially exhibit yellowing of leaves in the inner whorl, gradually spreading to the outer whorl of the crown. Stems of the affected palms become spongy and friable, and the conducting strands become destroyed. Virion properties and genome The virions are non-enveloped, filamentous particle about 1500–1700 nm in length. The genome is a positive-sense single-stranded RNA and consists of 16,080 nt with 11 open reading frames (ORFs) and a 11-nt-long 50 UTR and a 37-nt-long 30 UTR (KR349464) (Yu et al. 2015).

References Yu H, Qi S, Chang Z, Rong Q, Akinyemi IA, Wu Q (2015) Complete genome sequence of a novel Velarivirus infecting areca palm in China. Arch Virol 160(9):2367–2370

Argemone mexicana (Mexican prickly poppy) Family: Papaveraceae

Weed host

Bidens mottle virus Taxonomic position Genus: Potyvirus

(BiMoV)

Family: Potyviridae

BiMoV infection in plants of Argemone mexicana was reported from Florida, USA (Purcifull and Zitter 1973). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of BiMoV, refer to Bidens spp.

References Purcifull DE, Zitter TA (1973) A serological test for distinguishing bidens mottle and lettuce mosaic viruses. Fla Agric Exp Stations J (Series No. 5132):143–145

Argyranthemum frutescens (Marguerite daisy) Synonyms Chrysanthemum frutescens Family: Asteraceae

Ornamental

A

190

Argyranthemum frutescens (Marguerite daisy)

Chrysanthemum stunt viroid Taxonomic position Genus: Pospiviroid

(CSVd)

Family: Pospiviroidae

CSVd was reported on Argyranthemum frutescens (marguerite daisy) cv. Butterfly from Germany in 2000 (Menzel and Maiss 2000). The viroid infection in plants of A. frutescens was reported from France, Italy, and the UK (Marais et al. 2011; Torchetti et al. 2012; Harju et al. 2011). The viroid was detected from A. frutescens cv. Butterfly plants showing growth reduction, flower distortion, or leaf necrosis symptoms (Menzel and Maiss 2000) or from those showing yellow deformed leaves with terminal necrosis (Marais et al. 2011). The largely symptomless nature of CSVd in A. frutescens raises the possibility that contaminated plants may act as a reservoir for the future epidemics in chrysanthemum. The viroid is transmitted through foliar contact, by handling during cultivation, and by tools like cutting knives and scissors; and also transmitted mechanically through sap-inoculation. For more details of CSVd, refer to Chrysanthemum spp.

Chrysanthemum virus B Taxonomic position Genus: Carlavirus

(CVB)

Family: Betaflexiviridae

CVB infection in plants of Chrysanthemum frutescens was reported from Japan (Nakamura et al. 1994). The virus-infected marguerite daisy plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CVB, refer to Chrysanthemum spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Chrysanthemum frutescens was reported from Japan (Nakamura et al. 1994). The virus-infected marguerite daisy plants exhibit mosaic symptoms. The virus is transmitted by a number of aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of host plants. For more details of CMV, refer to Cucumis sativus.

Potato spindle tuber viroid Taxonomic position Genus: Pospiviroid

(PSTVd)

Family: Pospiviroidae

PSTVd incidence at low levels in plants of Argyranthemum frutescens was reported from the Czech Republic (Matousek et al. 2014). The viroid is mechanically sap-transmissible. The use of infected planting material is the primary mode of spread. For more details of PSTVd, refer to Solanum tuberosum.

Arisaema spp.

191

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Chrysanthemum frutescens was reported from Italy (Dellavalle et al. 1999). The virus-infected marguerite daisy plants show yellow or necrotic spotting and were stunted. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Dellavalle G, Ciuffo M, Roggero P, Lisa V, Minuto A, Minuto G, Rapetti S (1999) Tospoviruses in Delphinium sp., Gazania, Marguerite, Celery, Tragopogon porrifolius and Solanum rantonnetti in Liguria (Northern Italy). Informatore Fitopatologico 49:63–64 Harju VA, Skelton AL, Monger WA, Forde SMD, Daly M, Nixon T, Lawson R, Bennett S, Mumford RA (2011) A report of viruses, viroids and phytoplasmas found in ornamental plants from 1999–2007: findings from Central Science Laboratory, UK. Acta Hortic 901:223–229 Marais A, Faure C, Deogratias JM, Candresse T (2011) First report of Chrysanthemum stunt viroid in various cultivars of Argyranthemum frutescens in France. Aust Plant Dis Notes 95:1196 Matousek J, Piernikarczyk RJJ, Dedic P, Mertelik J, Uhlirova K, Duraisamy GS, Orctova L, Kloudova K, Ptacek J, Steger G (2014) Characterization of Potato spindle tuber viroid (PSTVd) incidence and new variants from ornamentals. Eur J Plant Pathol 138:93–101 Menzel W, Maiss E (2000) Detection of Chrysanthemum stunt viroid (CSVd) in cultivars of Argyranthemum frutescens by RT-PCR-ELISA. J Plant Dis Prot 107:548–552 Nakamura Y, Fujisawa I, Lee KH, Uematsu S (1994) Cucumber mosaic virus and Chrysanthemum virus B isolated from marguerite, Chrysanthemum frutescens L. Proc Kanto-Tosan Plant Protect Soc 41:177–179 Torchetti EM, Navarro B, Trisciuzzi VN, Nuccitelli L, Silletti MR, Di Serio F (2012) First report of Chrysanthemum stunt viroid in Argyranthemum frutescens in Italy. J Plant Pathol 94:451–454

Arisaema spp. Family: Araceae

Arisaema potyvirus 1

Ornamental

(ArV1)

Taxonomic position ArV1 is a tentative member of the genus Potyvirus and family Potyviridae. Geographical distribution ArV1 infection in plants of Arisaema spp. was reported from the Netherlands (Pham et al. 2008). Transmission The virus is presumed to be transmitted by aphid vectors. Virion properties and genome The virions are flexuous filaments. The genome consists of a single molecule of positive-sense singlestranded RNA. A partial genome sequence of 1331 nt is available (FJ546415) (Wylie et al. 2017).

A

192

Aristolochia spp.

Arisaema potyvirus 2

(ArV2)

Taxonomic position ArV2 is a tentative member of the genus Potyvirus and family Potyviridae. Geographical distribution ArV2 infection in plants of Arisaema spp. was reported from the Netherlands (Pham et al. unpublished FJ546415). Transmission The virus is presumed to be transmitted by aphid vectors. Virion properties and genome The virions are flexuous filaments. The genome consists of a single molecule of positive-sense singlestranded RNA. A partial genome sequence of 1014 nt is available (FJ546416) (Wylie et al. 2017).

Dasheen mosaic virus Taxonomic position Genus: Potyvirus

(DsMV)

Family: Potyviridae

DsMV infection in plants of Arisaema spp. was reported from Hawaii (Nelson 2008). The virus-infected plants exhibit systemic mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of DsMV, refer to Colocasia esculenta.

References Nelson SC (2008) Dasheen mosaic of edible and ornamental aroids. Plant Disease CTAHR. Available at: http://www. ctahr.hawaii.edu/oc/freepubs/pdf/PD-44.pdf Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Aristolochia spp. Family: Aristolochiaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Aristolochia zolligeriana was reported from Europe and Taiwan (Horvath et al. 1975; Chen et al. 2000). The virus-infected plants exhibit mosaic symptoms on the leaves. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Armoracia rusticana (Horseradish)

Grapevine fanleaf virus Taxonomic position Genus: Nepovirus

193

(GFLV)

A

Family: Secoviridae

GFLV infection in plants of Aristolochia clematitis was reported from Hungary (Horvath et al. 1994). The virus-infected plants exhibit symptoms of vein-clearing, yellow mosaic spots, and yellowing symptoms. The virus is transmitted by a nematode vector, Xiphinema index, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of GFLV, refer to Vitis vinifera.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Aristolochia clematitis was reported from Greece and Hungary (Chatzivassiliou et al. 2001; Cseh et al. 2013). The virus-infected plants exhibit yellow mosaic symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Chatzivassiliou EK, Boubourakas I, Drossos E, Eleftherohorinos I, Jenser G, Peters D, Katis NI (2001) Weeds in greenhouses and tobacco fields are differentially infected by Tomato spotted wilt virus and infested by its vector species. Plant Dis 85:40–46 Chen CC, Chang CA, Lin MJ, Fang HS (2000) Identification for the infection of Cucumber mosaic cucumovirus on Aristolochia zolligeriana. Plant Pathol Bull 9:29–34 Cseh E, Apro M, Bese G, Krizbai L, Boka K, Gaborjanyi R, Takacs AP (2013) Tomato spotted wilt virus (TSWV) in birthwort (Aristolochia clematitis L.) in Hungary. Acta Phytopathol Entomol Hung 48:33–38 Horvath J, Mamula D, Juretic N (1975) Some data concerning natural hosts of Cucumber mosaic virus in Hungary and Jugoslavia. Acta Bot Croat 34:9–16 Horvath J, Tobias I, Hunyadi K (1994) New natural herbaceous hosts of Grapevine fanleaf nepovirus. Kerteszeti Tudomany 26:31–32

Armoracia rusticana (Horseradish) Family: Brassicaceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Vegetable

(ArMV)

Family: Secoviridae

194

Armoracia rusticana (Horseradish)

ArMV infection in plants of Armoracia rusticana was reported from the UK and Czechoslovakia (Hickman and Varma 1968; Novak and Lanzova 1982). This virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Cauliflower mosaic virus Taxonomic position Genus: Caulimovirus

(CaMV)

Family: Caulimoviridae

CaMV infection in plants of Armoracia rusticana was reported from Illinois (Fletcher et al. 1984). The virus-infected horseradish plants exhibit vein-clearing or chlorotic mottling symptoms. The virus is transmitted by aphid vectors in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of CaMV, refer to Brassica oleracea var. botrytis.

Horseradish curly top virus Taxonomic position Genus: Curtovirus

(HCTV)

Family: Geminiviridae

Geographical distribution HCTV infection in plants of Armoracia rusticana was reported from Illinois, USA (Klute et al. 1996).

Symptoms and host(s) The virus-infected horseradish plants show either a general chlorosis or no symptoms. The virus has a narrow host range.

Transmission The virus is transmitted by the leafhopper vector, Circulifer tenellus, in a persistent (circulative, nonpropagative) manner. Experimentally shepherd’s purse (Capsella bursa-pastoris) and pennycress (Thlaspi arvense) have been shown to be susceptible to HCTV.

Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T=1, 22  38 nm with a single coat protein. Only a single isolate of HrCTV has been characterized. The genome of the virus consists of a single circular ssDNA of 3080 nt (U49907 = NC_002543). The genome encodes six genes; three in the virion-sense and three in complementary-sense orientation (Klute et al. 1996; Varsani et al. 2014; Zerbini et al. 2017). Oddly the isolate lacks the C3 gene that, for begomoviruses and curtoviruses, encodes the replication enhancer protein. Whether this is typical of the species, or just this isolate, is unclear. The expression and function of the genes have not been investigated (Klute et al. 1996; Briddon and Stenger 2011; Varsani et al. 2014).

Armoracia rusticana (Horseradish)

Horseradish latent virus Taxonomic position Genus: Caulimovirus

195

(HRLV)

Family: Caulimoviridae

Geographical distribution HRLV was first reported in plants of Armoracia rusticana from the USA and Denmark (Richins and Shepherd 1986). Symptoms and host(s) The virus-infected horseradish plants do not produce any conspicuous symptoms in American cultivars. The virus has narrow host range. Transmission The virus is transmitted by an aphid vector, Myzus persicae, in a semi-persistent manner. The virus is also transmitted by mechanical sap-inoculation. Virion properties and genome The virions are isometric, non-enveloped, 50 nm in diameter with no obvious surface structure. The genome is a single molecule of circular dsDNA which contains 7954 bp (JX429923 = NC_018858). The genome consists of six major open reading frames (Richins and Shepherd 1986; Hohn 2011).

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Armoracia rusticana was reported from New Zealand (Thomas and Procter 1973). The virus-infected horseradish plants do not exhibit any external symptoms. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Armoracia rusticana was reported from Brazil and the USA (Pound 1949; Fletcher et al. 1984; Horwitz et al. 1985; Eiras et al. 2007). The virus-infected horseradish plants exhibit symptoms of chlorotic mosaic or ring-spots on leaf blades, dark streaks on petioles, and rarely veinclearing. The virus is transmitted by aphid vectors, Myzus persicae, Brevicoryne brassicae, and other aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TuMV, refer to Brassica rapa.

A

196

Arracacia xanthorrhiza (Arracacha)

References Briddon RW, Stenger DC (2011) Curtovirus. Geminiviridae. In: The Springer index of Viruses. Springer, New York, pp 589–596. https://doi.org/10.1007/978-0-387-95919-1_82 Eiras M, Chaves ALR, Colariccio A, Chagas CM (2007) First report of Turnip mosaic virus in Horse radish in Brazil. Fitopatol Bras 32:165 Fletcher J, Schultz GA, Eastman CE (1984) Effect of mosaic viruses on infection of horse radish by Spiroplasma citri. Plant Dis 68:565–567 Hickman AJ, Varma A (1968) Viuses in horseradish. Plant Pathol 17:26–30 Hohn T (2011) Caulimovirus. Caulimoviridae. In: The Springer index of Viruses. Springer, New York, pp 271–277. https://doi.org/10.1007/978-0-387-95919-1_41 Horwitz DK, Fletcher J, D’Arcy CJ, Rhodes AM (1985) Turnip mosaic virus in the Illinois horseradish germ plasm collection. Plant Dis 69:246–248 Klute KA, Nadler SA, Stenger DC (1996) Horseradish curly top virus is a distinct subgroup II Geminivirus species with rep and C4 genes derived from a subgroup III ancestor. J Gen Virol 77:1369–1378 Novak JB, Lanzova J (1982) Arabis mosaic virus on rhubarb, cucumber and horseradish in Czechoslovakia. Sb UVTIZ Ochrana Rostlin 18:177–180 Pound GS (1949) The effect of air temperature on virus concentration and leaf morphology of mosaic-infected horseradish. J Agric Res 78:161–170 Richins RD, Shepherd RJ (1986) Horseradish latent virus, a new member of the Caulimovirus group. Phytopathology 76:749–754 Thomas W, Procter CH (1973) Tobacco ringspot virus in horseradish. N Z J Agric Res 16:233–237 Varsani A, Martin DP, Navas-Castillo J, Moriones E, Hernández-Zepeda C, Idris A, Murilo Zerbini F, Brown JK (2014) Revisiting the classification of curtoviruses based on genome-wide pairwise identity. Arch Virol 159(7):1873–1882 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Arracacia xanthorrhiza (Arracacha) Family: Apiaceae

Arracacha latent virus

Root vegetable

(ALV)

Taxonomic position ALV is a tentative member of the genus Carlavirus and family Betaflexiviridae. Geographical distribution ALV infection in plants of Arracacia xanthorrhiza was reported from the UK, South and Central America, Bolivia, and Peru (Brunt et al. 1996). Symptoms and host(s) The virus-infected Arracacha plants do not exhibit any symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is transmitted by mechanical sap-inoculation, and also through vegetative propagative plant material.

Arracacia xanthorrhiza (Arracacha)

197

Virion properties and genome The virions are filamentous, non-enveloped, and usually straight with a clear modal length of 650 nm (Brunt et al. 1996).

Arracacha mottle virus Taxonomic position Genus: Potyvirus

(AMoV)

Family: Potyviridae

Geographical distribution AMoV infection in plants of Arracacia xanthorrhiza was reported from Brazil (Orilio et al. 2009, 2013). Symptoms and host(s) The virus-infected arracacha plants exhibit mosaic mottling symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is mechanically saptransmissible. The virus is also transmitted through vegetative propagative plant material. Virion properties and genome The virions are non-enveloped, flexuous filaments rods of 730–740 nm long and 11–12 nm in diameter. The genome is a single molecule of positive-sense ssRNA of 9630 nt (DQ925486 = NC_018176) (Orilio et al. 2013).

Arracacha virus A Taxonomic position Genus: Nepovirus

(AVA)

Family: Secoviridae

Geographical distribution AVA infection was first reported in plants of Arracacia xanthorrhiza (arracacha) from the Peruvian Andes by Jones and Kenten (1978). The virus spreads in the South and Central American region and Peru (Adams et al. 2017). Symptoms and host(s) The virus-infected arracacha plants show a distinct yellow mosaic on young leaves. Transmission The virus is transmitted by nematode vectors in a non-persistent manner. The virus is readily transmissible by mechanical sap-inoculation, and has a wide host range, infecting species in at least ten dicotyledonous families. The virus is also transmissible through vegetative propagative plant material. The virus is not transmissible by contact between plants.

A

198

Arracacia xanthorrhiza (Arracacha)

Virion properties and genome The virions are isometric, non-enveloped of two types but similar in size 25–30 nm in diameter, and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is bipartite, linear positive-sense, single-stranded RNA (Sanfacon et al. 2009; Dunez and Le Gall 2011; Sanfacon 2015; Thompson et al. 2017); with RNA 1 of 7387 nt (KY569301), and RNA 2 of 3830 nt (KY569302).

Arracacha virus B Taxonomic position Genus: Cheravirus

(AVB)

Family: Secoviridae

Geographical distribution AVB was first reported in plants of Arracacia xanthorrhiza (arracacha) from the Peruvian Andes by Kenten and Jones (1979). The virus spreads in the South and Central American region, Bolivia, and Peru (Kenten and Jones 1979). Symptoms and host(s) The virus-infected arracacha plants do not exhibit any symptoms. Transmission The virus is transmitted by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants. The virus is also transmissible through vegetative propagative plant material. Virion properties and genome The virions are non-enveloped, 25–30 nm in diameter, and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is bipartite, linear positive-sense, single-stranded RNA. RNA1 consists of 7253 nt (JQ437415 = NC_020898) and RNA2 of 3486 nt (JQ581051 = NC_020897) (Sanfacon et al. 2009; Yoshikawa 2011; Adams et al. 2013; Sanfacon 2015; Le Gall et al. 2017; Thompson et al. 2017).

Arracacha virus V Taxonomic position Genus: Vitivirus

(AVV)

Family: Betaflexiviridae

Geographical distribution AVV infection in plants of Arracacia xanthorrhiza was reported from Brazil (Oliveira et al. 2017). Transmission The virus is mechanically sap-transmissible. The virus is primarily spread through the use of virusinfected vegetative planting material. Virion properties and genome The virions are helically constructed flexuous filaments, 750–800  12 nm, showing distinct crossbanding. The genome consists of a single molecule of positive sense ssRNA of 7398 nt in length (KY392781 = NC_034264) and contains five slightly overlapping ORFs which encode, in order, the

Arracacia xanthorrhiza (Arracacha)

199

replication-related proteins, a 19–20 K protein with unknown functions, an MP of the “30 K” superfamily type, the CP, and a small protein (10–14 K) with nucleotide binding properties. Coat protein subunit is of one type, and 18–21.5 kDa in size (Oliveira et al. 2017).

Bidens mosaic virus Taxonomic position Genus: Potyvirus

(BiMV)

Family: Potyviridae

BiMV infection in plants of Arracacia xanthorriza was reported from Brazil (Orilio et al. 2017). The virus-infected plants exhibit mottling symptoms. The virus is transmitted by aphid vectors in a nonpersistent manner, and also by mechanical sap-inoculation. For more details of BiMV, refer to Bidens spp.

Potato black ringspot virus Taxonomic position Genus: Nepovirus

(PBRSV)

Family: Secoviridae

PBRSV infection in plants of Arracacia xanthorrhiza was reported from Peru (Lizarraga et al. 1994). The virus-infected arracacha plants exhibit mosaic and leaf deformation symptoms. There is no known vector reported for this virus. The virus is transmitted by mechanical sap inoculation, and also by grafting. It is easily transmitted by contact between plants and through tubers. For more details of PBRSV, refer to Solanum tuberosum.

Potato virus S Taxonomic position Genus: Carlavirus

(PVS)

Family: Betaflexiviridae

PVS infection in plants of Arracacia xanthorrhiza was reported from Peru (De Souza et al. 2018). The virus is transmitted by aphid vectors, Myzus persicae and Macrosiphum euphorbiae, in a non-persistent manner, and also through mechanical sap-inoculation. For more details of PVS, refer to Solanum tuberosum.

References Adams IP, Glover R, Souza-Richards R, Bennett S, Hany U, Boonham N (2013) Complete genome sequence of Arracacha virus B: a novel Cheravirus. Arch Virol 158(4):909–913 Adams IP, Boonham N, Jones RAC (2017) First complete genome sequence of Arracacha virus A isolated from a 38-yearold sample from Peru. Genome Announc 5(18). pii: e00141-17. https://doi.org/10.1128/genomeA.00141-17 Brunt A, Crabtree K, Dallwitz M, Gibbs A, Watson L (1996) Viruses of plants: descriptions and lists from the VIDE database. 1484 pp. C.A.B.. International, UK De Souza J, Gamarra H, Muller G, Kreuze J (2018) First report of Potato virus S naturally infecting arracacha (Arracacia xanthorrhiza) in Peru. Plant Dis 102:460

A

200

Arrhenatherum elatius (Tall oat-grass)

Dunez J, Le Gall O (2011) Nepovirus. Comoviridae. In: The Springer index of Viruses. Springer, New York, pp 361–369. https://doi.org/10.1007/978-0-387-95919-1_55 Jones RAC, Kenten RH (1978) Arracacha virus A, a newly recognized virus infecting arracacha (Arracacia xanthorrhiza; Umbelliferae) in the Peruvian Andes. Ann Appl Biol 90:85–91 Kenten RH, Jones RAC (1979) Arracacha virus B, a second isometric virus infecting arracacha (Arracacia xanthorrhiza: Umbelliferae) in the Peruvian Andes. Ann Appl Biol 93:31–36 Le Gall O, Sanfacon H, Ikegami M, Iwanami T, Jones T, Karasev A, Lehto K, Wellink J, Wetzel T, Yoshikawa N (2017) Cheravirus and Sadwavirus: two unassigned genera of plant positive-sense single-stranded RNA viruses formerly considered atypical members of the genus Nepovirus (family Comoviridae). Arch Virol 152:1767–1774 Lizarraga C, Chuquillanqui C, Jayasinghe U (1994) A strain of PBRV (Potato black ringspot virus) isolated from arracacha (Arracacia xanthorrhiza). Fitopatología 29:144–149 Oliveira LM, Orilio AF, Inoue-Nagata AK, Nagata T, Blawid R (2017) A novel vitivirus-like sequence found in Arracacia xanthorrhiza plants by high throughput sequencing. Arch Virol 162:2141–2144 Orilio AF, Dusi AN, Madeira NR, Inoue-Nagata AK (2009) Characterization of a member of a new Potyvirus species infecting arracacha in Brazil. Arch Virol 154:181–185 Orilio AF, Lucinda N, Dusi AN, Nagata T, Inoue-Nagata AK (2013) Complete genome sequence of Arracacha mottle virus. Arch Virol 158(1):291–295 Orilio AF, Dusi AN, Madeira NR, Inoue-Nagata AK (2017) First report of Bidens mosaic virus in Arracacha (Arracacia xanthorriza) from Brazil. Plant Dis 101:262 Sanfacon H (2015) In: eLS (ed) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. Wiley, Chichester. http://www.els.net. https://doi.org/10.1002/9780470015902.a0000764.pub3 Sanfacon H, Wellink J, Le Gall O, Karasev A, van der Vlugt R, Wetzel T (2009) Secoviridae: a proposed family of plant viruses within the order Picornavirales that combines the families Sequiviridae and Comoviridae, the unassigned genera Cheravirus and Sadwavirus, and the proposed genus Torradovirus. Arch Virol 154(5):899–907 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J Gen Virol 98:529–531 Yoshikawa N (2011) Cheravirus. Sequiviridae. In: The Springer index of Viruses. Springer, New York, pp 1763–1768. https://doi.org/10.1007/978-0-387-95919-1_286

Arrhenatherum elatius (Tall oat-grass) Family: Poaceae

Weed host

Tall oatgrass mosaic virus Taxonomic position Genus: Tritimovirus

(TOgMV)

Family: Potyviridae

Geographical distribution TOgMV infection in plants of Arrhenatherum elatius was reported from the Czech Republic (Hassan et al. 2014). Symptoms and host(s) The virus-infected tall oat-grass plants exhibit mosaic symptoms. Transmission The virus is transmitted by mechanical sap-inoculation to A. elatius and a small number of other grasses, in which mosaic symptoms were induced; but the virus appears to have a narrow host range.

Artemisia spp. (A. annua; A. absinthium; A. vulgaris) (Wormwood)

201

Virion properties and genome The virions are non-enveloped, flexuous filaments, 700–720 nm long, and 12–15 nm in diameter. The genome is a monopartite, linear, positive-sense ssRNA of 9539 nt (KF260962 = NC_022745), with a 30 -poly(A) terminus (Lopez-Moya et al. 2009; French et al. 2011; Hassan et al. 2014; Wylie et al. 2017).

References French RC, Stenger DC, Tatineni S (2011) Tritimovirus. Potyviridae. In: The Springer index of Viruses. Springer, New York, pp 1445–1449. https://doi.org/10.1007/978-0-387-95919-1_237 Hassan M, Sirlova L, Vacke J (2014) Tall oatgrass mosaic virus (TOgMV): a novel member of the genus Tritimovirus infecting Arrhenatherum elatius. Arch Virol 159:1585–1592 Lopez-Moya JJ, Valli A, Garcia JA (2009) Potyviridae. In: Iacobellis NS, Collmer A, Hutcheson SW, Mansfield JW, Morris CE, Murillo J, Schaad D NW, Stead E, Surico G, M Ullrich (eds.). Encyclopaedia of life sciences (ELS). Kluwer Academic, Dordrecht, The Netherlands. https://doi.org/10.1002/9780470015902.a0000755.pub2 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Artemisia spp. (A. annua; A. absinthium; A. vulgaris) (Wormwood) Family: Asteraceae

Medicinal

Apple mosaic virus

(ApMV)

Taxonomic position Genus: Ilarvirus

Family: Bromoviridae

ApMV infection in plants of Artemisia vulgaris was reported from Turkey (Arli Sokmen et al. 2005). The virus-infected wormwood plants exhibit mild mosaic symptoms. No insect vector is known for this virus. The virus is mechanically sap-transmissible. The virus spreads by the use of infected vegetative planting material and also by graft inoculation (Paunovic et al. 2011). For more details of ApMV, refer to Malus domestica.

Artemisia virus A

(ArtVA)

Taxonomic position Genus: Sobemovirus

Family: Solemoviridae

Geographical distribution ArtVA infection in plants of Artemisia annua was reported from Switzerland (Ramel et al. 2013). Symptoms and host(s) The virus-infected wormwood plants exhibit dwarfing and chlorosis symptoms.

A

202

Artemisia spp. (A. annua; A. absinthium; A. vulgaris) (Wormwood)

Transmission The virus is transmitted by contact and the insect vector of the virus is not known. In the contaminated soil, healthy A. annua plant efficiently acquires the virus. The virus spreads through the use of infected planting material. Virion properties and genome The virions are isometric, non-enveloped, and 30 nm in diameter. The genome is polycistronic positivesense single-stranded RNA (ssRNA) of 4138 nt encoding 4 proteins (JN620802 = NC_017914). The genome consists of two overlapping ORFs, ORF2a, and ORF2b (Ramel et al. 2013; Somera et al. 2015).

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Artemisia dracunculus was reported from Pennsylvania (USA) and Bulgaria (Hausbeck et al. 1992; Dikova 2011). The virus-infected wormwood plants exhibit chlorosis symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Artemisia annua was reported from China (Ding et al. 2008). The virusinfected wormwood plants exhibit symptoms of curling of the leaves and shoots. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. The virus is transmissible by grafting but not by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

References Arli Sokmen M, Kutluk Yilmaz ND, Mennan H, Sevik MA (2005) Natural weed hosts of Apple mosaic virus in hazelnut orchards in Turkey. J Plant Pathol 87:239–242 Dikova B (2011) Tomato spotted wilt virus on some medicinal and essential oil-bearing plants in Bulgaria. Bulg J Agric Sci 17:306–313 Ding M, Yue N, Zhang ZK, Zhao ZW (2008) First report of Tomato yellow leaf curl virus in Artemisia annua in China. J Plant Pathol 90(3):589 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among greenhouse ornamentals in Pennsylvania. Plant Dis 76:795–800 Paunovic S, Pasquini G, Barba M (2011) Apple mosaic virus in stone fruits. In: Hadidi A, Barba M, Candresse T, Jelkmann W (eds) Virus and virus-like diseases of pome and stone fruits. APS Press, St. Paul, pp 91–95 Ramel M-E, Gugerli P, Besse S, Simonnet X, Balmelli C (2013) Molecular characterization of Artemisia virus A, a new Sobemovirus isolated from Artemisia annua. Arch Virol 158:458–466 Somera M, Sarmiento C, Truve E (2015) Overview on Sobemoviruses and a proposal for the creation of the family Sobemoviridae. Viruses 7:3076–3115

Arum spp. (Arum lilies)

203

Arthropodium cirratum (Renga lily) Family: Asparagaceae

Ornamental

Cucumber mosaic virus

(CMV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV infection in plants of Arthropodium cirratum was reported from New Zealand (Fletcher 2001). The virus-infected renga lily plants exhibit symptoms of leaves with diffuse mottling on stunted plants. The virus is transmitted by a number of aphid vectors in the non-persistent manner, and also by mechanical sap-inoculation to a large number of host plants. For more details of CMV, refer to Cucumis sativus.

References Fletcher JD (2001) New hosts of Alfalfa mosaic virus, Cucumber mosaic virus, Potato virus Y, Soybean dwarf virus, and Tomato spotted wilt virus in New Zealand. N Z J Crop Hortic Sci 29:213–217

Arum spp. (Arum lilies) Family: Araceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

Family: Secoviridae

ArMV infection in plants of Arum spp. was reported from Lithuania and Iran (Ghotbi and Shahraeen 2005; Samuitiene et al. 2008). The virus-infected arum lily plants were smaller than normal. Leaves show yellow spots and streaks. The virus is transmitted by the nematode vectors, Xiphinema diversicaudatum and X. coxi, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

A

204

Asclepias spp.

TSV infection in plants of Arum orientale was reported from Iran (Ghotbi 2006). The virus-infected arum lily plants exhibit symptoms of small necrotic lesions on the leaves. The virus is transmitted by thrips vectors. The virus present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible. The virus is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Arum spp. was reported from California, Hawaii, Italy, and France (Tompkins and Severin 1950; Cho et al. 1987; Parrella et al. 2003). The virus-infected arum lily plants exhibit symptoms of chlorotic to whitish streaks and interveinal circular lesions that may become necrotic (Tompkins and Severin 1950). The virus is transmitted by thrips vectors in a persistent, propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting, but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Cho JJ, Mau RFL, Mitchell WC, Gonsalves D, Yudin LS (1987) Host list of plants susceptible to Tomato spotted wilt virus (TSWV), Research Extension Series; RES-078. University of Hawaii, Honolulu 10 p Ghotbi T (2006) First report on incidence of Tobacco streak virus (TSV) on ornamental plants in Iran. Iran J Plant Pathol 42:159–160 Ghotbi T, Shahraeen N (2005) First report on incidence of Arabis mosaic virus (ArMV, Nepovirus) on ornamental plants in Iran. Iran J Plant Pathol 41(2):305–306 Parrella G, Gognalons P, Gebre-Selassi K, Vovlas C, Marchoux G (2003) An update of the host range of Tomato spotted wilt virus. J Plant Pathol 85:227–264 Samuitiene M, Navalinskiene M, Jackeviciene E (2008) Arabis mosaic virus on ornamental plants. Biologia 54:264–268 Tompkins CM, Severin HHP (1950) Spotted wilt of white, yellow, and pink callas. Hilgardia 20:207–232

Asclepias spp. Family: Apocynaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Ornamental

(AMV)

Family: Bromoviridae

AMV was isolated from naturally infected plants of Asclepias syriaca in Hungary and the USA (Kazinczi et al. 2002). The virus-infected plants exhibit chlorotic spotting or bright yellow mosaic symptoms. The virus is transmitted by a large number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Asclepias spp.

Arabis mosaic virus Taxonomic position Genus: Nepovirus

205

(ArMV)

Family: Secoviridae

ArMV infection in plants of Asclepias syriaca was reported from Italy (Bellardi and Rubies-Autonell 1999). The virus-infected plants exhibit symptoms of chlorotic mosaic, line patterns, and yellow spots or rings. This virus is transmitted by nematode vectors (Xiphinema spp.) in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Asclepias asymptomatic virus

(AsAV)

Taxonomic position AsAV is a tentative member of the genus Tymovirus and family Tymoviridae

Geographical distribution AsAV infection in plants of Asclepias viridis was reported from Oklahoma (USA) (Min et al. 2012).

Symptoms and host(s) The virus-infected Asclepias viridis plants do not exhibit any symptoms.

Transmission The virus is transmitted by beetle vectors, Tetraopes spp. The virus is mechanically sap-transmissible to Nicotiana benthamiana and develops chlorosis associated with mottling symptoms (Min et al. 2012).

Virion properties and genome The virions are isometric particles of c.28–30 nm. The full 6175 nt single-stranded RNA genome sequence is available (HQ425778 = NC_015523).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Asclepias spp. was detected in the USA, Hungary, Yugoslavia, and Israel (Horvath et al. 1983; Salamon 1986; Rist and Lorbeer 1989; Salamon et al. 1989; Gera 1994; Samuitiene and Navalinskiene 2008). The virus-infected plants show leaves with mosaic pattern of yellow patches or mottled yellow or pale green with normal leaf tissue, vein-clearing, line pattern, leaf curling, or miniaturization of leaf blades. The virus is transmitted by an aphid vector Aphis gossypii in a non-persistent manner, and also by mechanical sap-inoculation to a very wide host range (Ferreira and Boley 1992). For more details of CMV, refer to Cucumis sativus.

A

206

Asparagus officinalis (Asparagus)

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV occurs wherever Asclepias spp. are grown (Salomone et al. 2003). The virus-infected plants exhibit yellowing, mild mosaic, necrosis, and malformation of leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Bellardi MG, Rubies-Autonell C (1999) Occurrence of Arabis mosaic Nepovirus (ArMV) on Asclepias syriaca L. Phytopathol Mediterr 38:40–42 Ferreira SA, Boley RA (1992) Cucumber mosaic virus. Crop knowledge master. University of Hawaii, CTAHR, Manoa Gera A (1994) The natural occurrence of Cucumber mosaic virus in ornamentals in Israel. Acta Hortic 377:99–106 Horvath J, Mamula D, Salamon P (1983) Susceptibility of Asclepias syriaca L. (Asclepiadaceae) to Cucumber mosaic virus and its role in the virus epidemiology. Novenyvedelem 19:352–353 Kazinczi G, Horvath J, Lesemann DE (2002) Perennial plants as new natural hosts of three viruses. Z Pflanzenkrankh Pflanzenschutz 109:301–310 Min B-E, Feldman TS, Ali A, Wiley G, Muthukumar V, Roe BA, Roossinck M, Melcher U, Palmer MW, Nelson RS (2012) Molecular characterization, ecology, and epidemiology of a novel Tymovirus in Asclepias viridis from Oklahoma. Phytopathology 102:166–176 Rist DL, Lorbeer JW (1989) Occurrence and over wintering of Cucumber mosaic virus and Broad bean wilt virus in weeds grown near lettuce fields in New York. Phytopathology 79:65–69 Salamon P (1986) Asclepias syriaca L., a reservoir host of Cucumber mosaic virus in Hungary. Kertgazdasag 18:45–59 Salamon P, Horvath J, Mamula D, Juretic N, Hunyadi K (1989) Asclepias syriaca L. (common milkweed), a new natural host of Cucumber mosaic virus in Hungary and Yugoslavia. Acta Phytopathol Entomol Hung 24:363–373 Salomone A, Masenga V, Minuto G, Parodi C, Roggero P (2003) First report of Tomato spotted wilt virus (Tospovirus, Bunyaviridae) infecting Euphorbia eritrea and Asclepias curassavica in Liguria, Italy. Plant Pathol 52:806 Samuitiene M, Navalinskiene M (2008) Occurrence of Cucumber mosaic cucumovirus on ornamental plants in Lithuania. Zemdirbyste-Agriculture 95:135–143

Asparagus officinalis (Asparagus) Family: Liliaceae

Asparagus virus 1 Taxonomic position Genus: Potyvirus

Leafy vegetable

(AV-1)

Family: Potyviridae

Geographical distribution AV-1 infection in plants of Asparagus officinalis was first reported in 1960 (Hein 1960). The virus spreads wherever asparagus is grown commercially including the Eastern Asian region, the Eurasian region, the North American region, Germany, Italy, Japan, and the UK (Fujisawa et al. 1983a; Davis and Garrison 1984; Howell and Mink 1985; Falloon et al. 1986; Montasser and Davis 1987; Bertaccini

Asparagus officinalis (Asparagus)

207

et al. 1990; Evans 1991; Rafael et al. 1994; Tomassoli et al. 2007; Tiberini et al. 2008; Tomassoli et al. 2012; Tiberini et al. 2014; Li et al. 2017).

A Symptoms and host(s) The virus-infected asparagus plants do not produce any external symptoms. Asparagus is the only known natural host of this virus. Transmission The virus is transmitted by aphid vectors, Aphis craccivora and Myzus persicae, in a non-persistent manner. The virus is transmitted by mechanical sap-inoculation to a number of host species. The virus is not transmissible by contact between plants, not by seed, and not by pollen (Tomassoli et al. 2009). Virion properties and genome The virions are non-enveloped, flexuous filaments, 740 nm in length, and 13 nm in width. The genome consists of a single molecule of linear, positive-sense, single-stranded RNA of 9741 nt (KJ830760 = NC_025821). The RNA genome contains one large open reading frame (ORF) that is expressed as a single polyprotein about 350 kDa in size (Howell and Mink 1985; Tiberini et al. 2008; Tomassoli et al. 2008; Blockus et al. 2015; Wylie et al. 2017).

Asparagus virus 2 Taxonomic position Genus: Ilarvirus

(AV-2)

Family: Bromoviridae

Geographical distribution AV-2 infection in plants of Asparagus officinalis was reported from the USA, Germany, the Netherlands, England, Canada, and Japan (Fujisawa et al. 1983b; Hartung et al. 1985; Phillips and Brunt 1985; Falloon et al. 1986; Montasser and Davis 1987; Bertaccini et al. 1988, 1990; Evans 1991; Wolyn and Stobbs 1991; Rafael et al. 1994; Jaspers and Falloon 1996, 1999; Roose et al. 2002; Tomassoli et al. 2007, 2012). Symptoms and host(s) The virus-infected asparagus plants do not produce any external symptoms (Jaspers 1996; Jaspers et al. 1999). Asparagus is the only known natural host of this virus. Transmission The virus is transmitted by thrips vectors, the virus present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is mechanically sap-transmissible. Experimentally the virus-infected 116 of 169 species in 27 of 34 families of dicotyledonous plants. The virus also infected two of three species from three families of monocotyledonous plants (Mink and Uyeda 1977). The virus is transmitted mechanically on harvest knife blades, mowers, cultivation equipment, or any other activity that moves plant sap from one plant to another. The virus is seed-transmitted up to 60% in asparagus (Evans and Stephens 1988; Jaspers and Pearson 1997; Kawamura et al. 2014; Jaspers et al. 2015).

208

Asparagus officinalis (Asparagus)

Virion properties and genome The virus particles sediment as three major components known as NP1, NP2, and NP3 which are quasiisometric particles with modal diameters of 32 nm, 28 nm, and 26 nm, respectively (Uyeda and Mink 1981; Giunchedi et al. 1987). The genome consists of linear single-stranded RNA distributed among three RNA species: RNA1, 3431 nt (EU919666 = NC_011808); RNA2, 2916 nt (EU919667 = NC_011809); and RNA3, 2307 nt (X86352 = NC_011807) (Rafael-Martin and Rivera-Bustamante 1999; Scott and Zimmerman 2009; Scott 2011a, b).

Asparagus virus 3 Taxonomic position Genus: Potexvirus

(AV-3)

Family: Alphaflexiviridae

Geographical distribution AV-3 infection in plants of Asparagus officinalis was reported from Japan by Fujisawa (1986). The virus spreads in the Eastern Asian region and Japan (Bertaccini et al. 1990). Symptoms and host(s) The virus-infected asparagus plants exhibit symptoms of faint leaf chlorosis on young leaves. Transmission The virus is transmitted by means not involving a vector. The virus is transmitted by mechanical sapinoculation. The virus is not transmitted by seed. Virion properties and genome The virions are flexuous filaments with a clear modal length of 580 nm and 13 nm wide. The genome is positive-sense single-stranded RNA of 6937 nt (AB304848 = NC_010416) (Hashimoto et al. 2008) and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type, 18–27 kDa in size (Adams et al. 2004).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Asparagus officinalis was reported from Germany and Britain (Weissenfels and Schmelzer 1976; Phillips and Brunt 1985; Kegler et al. 1991; Bandte et al. 2008). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Tobacco streak virus Synonyms Asparagus stunt virus

(TSV)

Asparagus officinalis (Asparagus)

Taxonomic position Genus: Ilarvirus

209

Family: Bromoviridae

TSV infection in plants of Asparagus officinalis was reported from Denmark, Mexico, and California (USA) (Paludan 1964; Mink and Uyeda 1977; Uyeda 1978; Falloon et al. 1986; Wolyn and Stobbs 1991; Rafael et al. 1994). The virus was previously referred to as Asparagus stunt virus (Brunt and Paludan 1970). The virus-infected asparagus plants are stunted and show chlorotic or brown streaks on stems and browning of needles. The virus is transmitted by thrips vectors (Frankliniella occidentalis and Thrips tabaci), the virus present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is not seed-transmissible in asparagus and not by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Bandte M, Grubits E, von Bargen S, Rabenstein F, Weber D, Uwihs F, Buttner C (2008) Field study on the occurrence of virus infections in asparagus (Asparagus officinalis L.) in North Germany. “Erna¨hren uns in der Zukunft Energiepflanzen”, 63rd ALVA-Tagung in Raumberg-Gumpenstein, Austria, pp 97–99 Bertaccini A, Macri S, Poggi Pollini C (1988) Diagnosis of Asparagus virus 2 with ELISA method. Inf Fitopat 38:39–41 Bertaccini A, Giunchedi L, Poggi Pollini C (1990) Survey on asparagus diseases in Italy. Acta Hortic 271:279–283 Blockus S, Lesker T, Maiss E (2015) Complete genome sequences of two biologically distinct isolates of Asparagus virus 1. Arch Virol 160(2):569–572 Brunt AA, Paludan N (1970) The serological relationship between “Asparagus stunt virus” and Tobacco streak virus. Phytopathol Z 69:277–282 Davis RF, Garrison S (1984) First report of a virus present in asparagus in New Jersey. Plant Dis 68:628 Evans TA (1991) Occurrence of Asparagus virus I and Asparagus virus II on Asparagus in Delaware. Plant Dis 75:1075 Evans TA, Stephens CT (1988) Association of Asparagus virus II with pollen from infected asparagus (Asparagus officinalis). Plant Dis 72:195–198 Falloon PG, Falloon LM, Grogan RG (1986) Survey of California asparagus for Asparagus virus I, Asparagus virus II and Tobacco streak virus. Plant Dis 70:103–105 Fujisawa I (1986) Asparagus virus III: a new member of Potexvirus from asparagus. Ann Phytopathol Soc Jap 52:193–200 Fujisawa I, Goto T, Tsuchizaki T, Iizuka N (1983a) Host range and some properties of Asparagus virus I isolated from Asparagus officinalis in Japan. Ann Phytopathol Soc Jpn 49:299–307 Fujisawa I, Goto T, Tsuchizaki T (1983b) Some properties of Asparagus virus 2 isolated from Asparagus officinalis in Japan. Ann Phytopathol Soc Jpn 49:683–688 Giunchedi L, Poggi Pollini C, Bertaccini A, Marani F (1987) Purification and some properties of an Italian asparagus virus 2 isolate. Phytopathol Mediterr 26:117–120 Hartung AC, Evans TA, Stephens CT (1985) Occurrence of Asparagus virus II in commercial asparagus fields in Michigan. Plant Dis 69:501–504 Hashimoto M, Ozeki J, Komatsu K, Senshu H, Kagiwada S, Mori T, Yamaji Y, Namba S (2008) Complete nucleotide sequence of Asparagus virus 3. Arch Virol 153:219–221 Hein A (1960) On the occurrence of a virosis of Asparagus. Z PflKrankh 67:217–219 Howell WE, Mink GI (1985) Properties of Asparagus virus 1 isolated from Washington State asparagus. Plant Dis 69:1044–1046 Jaspers MV (1996) Effect of Asparagus virus 2 on yield of Asparagus officinalis. Acta Hortic 415:383–386 Jaspers MV, Falloon PG (1996) Survey of asparagus in New Zealand for Asparagus virus 2. Acta Hortic 415:301–307 Jaspers MV, Falloon PG (1999) Asparagus virus 2: a contributing factor in asparagus decline. Acta Hortic 479:263–270 Jaspers MV, Pearson MN (1997) Transmission of Asparagus virus 2 in an asparagus crop. In: Proceedings of the 50th New Zealand plant protection conference, pp 84–88 Jaspers MV, Falloon PG, Pearson MN (1999) Long-term effects of Asparagus virus 2 infection on growth and productivity in asparagus. Ann Appl Biol 135:379–384

A

210

Asplenium scolopendrium (Hart’s-tongue fern)

Jaspers MV, Falloon PG, Pearson MN (2015) Seed and pollen transmission of Asparagus virus 2. Eur J Plant Pathol 142:173–183 Kawamura R, Shimura H, Mochizuki T, Ohki ST, Masuta C (2014) Pollen transmission of Asparagus virus 2 (AV-2) may facilitate mixed infection by two AV-2 isolates in asparagus plants. Phytopathology 104:1001–1006 Kegler H, Schmidt HB, Wolterstorff B, Reinhardt I, Weber I, Proll E (1991) Spread of viruses in asparagus fields. Arch Phytopathol Plant Prot 27:251–258 Li MJ, Ke SY, Lin C, Mao ZC, Liu ZJ, Anane F, Zhao MF, Wen GS (2017) First report of Asparagus virus 1 on asparagus (Asparagus officinalis) in China. Plant Dis 101:844 Mink GI, Uyeda I (1977) Three mechanically-transmissible viruses isolated from asparagus in Washington. Plant Dis Reptr 61:398–401 Montasser MS, Davis RF (1987) Survey for Asparagus viruses I and II in New Jersey. Plant Dis 71:497–499 Paludan N (1964) Virus disease of Asparagus officinalis. Manedsovers Plantesygd 407:11–16 Phillips S, Brunt AA (1985) Occurrence of Cucumber mosaic virus and Asparagus virus II in asparagus (Asparagus officinalis L. var. officinalis) in Britain. Plant Pathol 34:440–442 Rafael-Martin M, Rivera-Bustamante RF (1999) Molecular characterization of the RNA3 of Asparagus virus 2. Arch Virol:185–192 Rafael M, Becerra A, Rivera-Bustamante R (1994) First report of viruses in asparagus plantings in Mexico. Plant Dis 78:1123 Roose ML, Stone NK, Mathews DM, Dodds JA (2002) RT-PCR detection of Asparagus 2 Ilarvirus. Acta Hortic 589:357–363 Scott SW (2011a) Bromoviridae and allies. In: Encyclopedia of life sciences (ELS). Wiley, Chichester. https://doi.org/ 10.1002/9780470015902 Scott SW (2011b) Ilarvirus. Bromoviridae. The Springer index of Viruses. Springer, New York, pp 87–194. https://doi. org/10.1007/978-0-387-95919-1_27 Scott SW, Zimmerman MT (2009) The nucleotide sequences of the RNA 1 and RNA 2 of Asparagus virus 2 show a close relationship to Citrus variegation virus. Arch Virol 154(4):719–722 Tiberini A, Tomassoli L, Vetten HJ (2008) Asparagus virus 1: biological, serological and molecular properties of a virus of the genus Potyvirus. Abstract IWGLVV conference, Ljubljana, p 29 Tiberini A, Zaccaria A, Tomassoli L (2014) Incidence and genetic variability of Asparagus virus 1 in naturally infected asparagus. J Plant Pathol 96:177–182 Tomassoli L, Zaccaria A, Valentino D, Tamietti G (2007) Preliminary investigation on asparagus diseases in Sicily. J Plant Pathol 89(3):S63 Tomassoli L, Tiberini A, Zaccaria A, Vetten JH (2008) Molecular and biological studies of Asparagus virus 1 (genus Potyvirus). J Plant Pathol 90(Suppl 2):437 Tomassoli L, Tiberini A, Zaccaria A, Falavigna A (2009) Prevalence of Asparagus virus 1 in asparagus commercial crops. Abstract XII International Asparagus Symposium, Lima Tomassoli L, Tiberini A, Vetten HJ (2012) Viruses of asparagus. Adv Virus Res 84:349 Uyeda I (1978) Identification, characterization, and incidence of viruses isolated from asparagus. PhD thesis, Washington State University, Pullman, 115 p Uyeda I, Mink GI (1981) Properties of Asparagus virus II, a new member of the Ilarvirus group. Phytopathology 71:1264–1269 Weissenfels M, Schmelzer K (1976) Untersuchungen uber das Schadausmass durch Viren am Spargel (Asparagus officinalis L.). Arch Phytopathol Pflanzenschutz 12:67–73 Wolyn DJ, Stobbs LW (1991) Distribution of Asparagus virus 2 and Tobacco streak virus in asparagus plantings in Southern Ontario. Plant Dis 75:430 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Asplenium scolopendrium (Hart’s-tongue fern) Synonyms Phyllitis scolopendrium Family: Aspleniaceae

Ornamental

Aster spp.

211

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Asplenium nidus-avis was reported from Spain (Lavina and Battle 1994). The virus-infected plants exhibit symptoms of ringspots and numerous necrotic lesions. The virus is transmitted by thrips vectors in a persistent, propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

Infection of Asplenium scolopendrium by TSWV has been reported from Iran (Ghotbi and Eskandari 2010; Ghotbi and Shahraeen 2012). The virus-infected plants exhibit symptoms of necrotic leaf spots and chlorosis. The virus is transmitted by thrips vectors in a persistent, propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting, but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Ghotbi T, Eskandari M (2010) First report of incidence Tomato spotted wilt virus (TSWV) on Asplenium scolopendrium in Iran. 19th Iranian Plant Protection Congress. Tehran, Iran. p 723 Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses affecting propagated ornamentals in Northern Iran. Int Res J Microbiol 3:373–381 Lavina A, Battle A (1994) First report of Impatiens necrotic spot virus in Asplenium nidus-avis in Spain. Plant Dis 78:316

Aster spp. Family: Asteraceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Aster yomena was reported from Korea, the Netherlands, and Israel (Loebenstein et al. 1995; Choi et al. 1990; Chen 2003). The virus-infected plants show mottle and severe mosaic symptoms. The virus is transmitted by a number of aphid vectors such as Myzus persicae and Aphis gossypii in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

A

212

Papaya leaf curl virus Taxonomic position Genus: Begomovirus

Aster spp.

(PaLCuV)

Family: Geminiviridae

PaLCuV infection in plants of Aster alpinus was reported from India (Snehi et al. 2013). The virusinfected plants exhibit yellow vein symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not mechanically sap-transmissible. For more details of PaLCuV, refer to Carica papaya.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in plants of Aster spp. was reported from Japan (Komuro et al. 1971). The virus-infected plants exhibit symptoms of ringspots, wrinkled curly leaves, thickened chlorotic rootlets, deformed flowers, and retarded growth. The virus is easily transmitted by mechanical means to a wide host range. Under field conditions, the virus is transmitted by nematodes, Trichodorus minor. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Aster spp. was reported from commercial nurseries in the USA (Lockhart et al. 2002). The virus-infected plants exhibit stunting and systemic blotching symptoms. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection of Aster spp. has been reported in multiple countries (Cho et al. 1987; Gera and Cohen 1995; Parrella et al. 2003). The virus-infected plants exhibit necrotic line patterns, ringspots on leaves, malformation, and flower distortion and may be stunted and exhibit systemic chlorosis. The necrotic lines persist as long as the affected leaves live while other symptoms may disappear. Symptoms are more severe in cool weather (Marchoux et al. 1991). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Astilbe spp.

213

References Chen YK (2003) Occurrence of Cucumber mosaic virus in ornamental plants and perspectives of transgenic control. PhD thesis, Wageningen University, The Netherlands. p 144 Cho JJ, Mau RFL, Mitchell WC, Gonsalves D, Yudin LS (1987) Host list of plants susceptible to Tomato spotted wilt virus (TSWV). Research Extension Series; RES-078. University of Hawaii, Honolulu. 10 p Choi JK, Kwon SB, Lee SY, Park WM (1990) Some properties of two isolates of Cucumber mosaic virus isolated from Aster yomena Makino and Commelina communis L. Korean J Plant Pathol 6:138–143 Gera A, Cohen J (1995) Aster. In: Loebenstein G, Lawson RH, Brunt AA (eds) Virus and virus-like diseases of bulb and flower crops. Wiley, Chichester Komuro Y, Yoshino M, Ichinoche M (1971) Tobacco rattle virus from aster showing ringspot syndrome and its transmission by Trichodorous minor, Colbran. Rev Plant Prot Res 4:130–132 Lockhart BEL, Fetzer J, Westendrop J (2002) Previously unreported viral diseases of Aster, Heuchera, Lobelia, Pulmonaria and Physostegia in the USA. Acta Hortic 568:221–224 Loebenstein G, Lawson RH, Brunt AA (1995) Virus and virus-like diseases of bulb and flowering crops. Wiley, Chichester, p 543 Marchoux G, Gebre-Selassie K, Villevieille M (1991) Detection of Tomato spotted wilt virus and transmission by Frankliniella occidentalis in France. Plant Pathol 40:347–351 Parrella G, Gognalons P, Gebre-Selassi K, Vovlas C, Marchoux G (2003) An update of the host range of Tomato spotted wilt virus. J Plant Pathol 85:227–264 Snehi SK, Srivastava A, Kumar S, Raj SK (2013) Molecular detection and identification of begomoviruses affecting important ornamental plants in India. Indian J Virol 24:142

Astilbe spp. Family: Saxifragaceae

Ornamental

Strawberry latent ringspot virus Taxonomic position Genus: Unassigned

(SLRSV)

Family: Secoviridae

SLRSV infection in plants of Astilbe x arendsii was reported from Finland (Bremer 1985). The virus is transmitted by nematode vector (Xiphinema spp.) and also by mechanical sap-inoculation. For more details of SLRSV, refer to Fragaria spp.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Astilbe chinensis was reported from Minnesota (USA) (Lockhart 2005). The virus-infected plants exhibit systemic chlorosis symptoms. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

A

214

Astragalus sinicus (Milk vetch)

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Astilbe spp. was reported from Ohio (USA) (Fisher 2013). The virus-infected plants exhibit necrosis and leaf mottle symptoms. The virus is transmitted by thrips vectors: the virus present in/on the pollen, entering the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato black ring virus Taxonomic position Genus: Nepovirus

(TBRV)

Family: Secoviridae

TBRV infection in plants of Astilbe x arendsii was reported from Finland (Tapio 1985). The virus is transmitted by the nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TBRV, refer to Solanum lycopersicum.

References Bremer K (1985) Strawberry latent ringspot virus in ornamental plants in Finland. Ann Agric Fenniae 24:101–102 Fisher JR (2013) Identification of two Tobacco streak virus capsid protein variants associated with leaf mottle and necrosis symptoms on Astilbe. Plant Health Prog. https://doi.org/10.1094/PHP-2013-0729-02-BR Online Lockhart BEL (2005) Three previously unrecorded viral diseases of Astilbe, Fuschia, and Thermopsis species in Minnesota. Plant Dis 89:775 Tapio E (1985) The appearance of soil-borne viruses in Finnish plant nurseries II. J Agric Sci Finl 57:167–181

Astragalus sinicus (Milk vetch) Family: Fabaceae

Milk vetch dwarf virus Taxonomic position Genus: Nanovirus

Medicinal

(MDV)

Family: Nanoviridae

Geographical distribution MDV infection was first reported in plants of Astragalus sinicus from Japan by Matsuura (1953). The virus spreads in Japan (Sano et al. 1993, 1998). Symptoms and host(s) The virus-infected milk vetch plants exhibit yellow dwarf and leaf rolling symptoms.

Asystasia gangetica (Chinese violet)

215

Transmission The virus is transmitted by aphid vectors, Aphis craccivora, A. gossypii, Acyrthosiphon pisum, and Acyrthosiphon (Aulacorthum) solani in a persistent but non-propagative manner. The virus is retained when the vector moults and does not multiply in a vector. The virus is not transmitted by mechanical inoculation. Under experimental conditions, the susceptible host species are found in the families Chenopodiaceae, Leguminosae-Caesalpinioideae, Leguminosae-Papilionoideae, and Solanaceae. The virus is transmitted by grafting. Virion properties and genome The virions are non-enveloped and spherical of about 18 nm in diameter with icosahedral symmetry. The genome consists of eight different positive-sense ssDNA components: DNA-C, 990 nt (AB000923); DNA-U1, 989 nt (AB000924); DNA-N, 977 nt (AB000925); DNA-U2, 981 nt (AB000926); DNA-M, 985 nt (AB000927); DNA-S, 997 nt (AB009046); DNA-R, 1001 nt (AB027511); and DNA-U4, 991 nt (AB255373) (NC_003641-NC_003646, NC_003648, NC_023626). Each ssDNA segment has a common stem-loop region and is encapsidated in a separate particle (Sano et al. 1993, 1998).

References Matsuura Y (1953) Studies on the dwarf disease of milk vetch (Astragalus sinicus). Ann Phytopathol Soc Jpn 17:65–68 Sano Y, Isogai M, Satoh S, Kojima M (1993) Small virus-like particles containing single-stranded DNAs associated with milk-vetch dwarf disease in Japan. 6th international congress of plant pathology, Montreal. Abstract no. 17.1.27. p 305 Sano Y, Wada M, Hashimoto Y, Matsumoto T, Kojima M (1998) Sequences of ten circular ssDNA components associated with the Milk vetch dwarf virus genome. J Gen Virol 79:3111–3118

Asystasia gangetica (Chinese violet) Family: Acanthaceae

Ornamental

Asystasia mosaic Madagascar virus Taxonomic position Genus: Begomovirus

(AsMMV)

Family: Geminiviridae

Geographical distribution AsMMV infection in plants of Asystasia gangetica was reported from Madagascar (De Bruyn et al. 2015). Symptoms and host(s) The virus-infected Chinese violet plants show bright yellow mosaic symptoms. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner.

A

216

Asystasia gangetica (Chinese violet)

Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2738 nt (KP663485 = NC_026514) and DNA-B of 2666 nt (KP663484 = NC_026513) (Briddon 2001; Brown et al. 2015; De Bruyn et al. 2015; Zerbini et al. 2017).

West African Asystasia virus 1

(WAAV1)

Synonyms Asystasia begomovirus 1 Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution WAAV1-infected plants of Asystasia gangetica were reported from West Africa (Wyant et al. 2015). Symptoms and host(s) The virus-infected Chinese violet plants exhibit vein yellowing, mosaic, and leaf curling symptoms. Transmission WAAV1 (possibly in combination with WAAV2) has been shown to be transmitted by a specific Bemisia tabaci “type” which transmitted other begomoviruses poorly (Bedford et al. 1992). In common with all other begomoviruses, WAAV1 is likely transmitted by the whitefly in a circulative, nonpropagative manner. Experimentally WAAV1 has been shown to infect Nicotiana benthamiana (exhibiting stunting) and Datura stramonium (exhibiting leaf curling and stunting) by biolistic inoculation of viral DNA. Virion properties and genome The structure of the virions of WAAV1 has not been investigated. In common with all geminiviruses, the virions of WAAV1 are likely geminate (twinned icosahedra). The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A contains 2727 nt (JF694484 = NC_016575) and DNA-B of 2661 nt (JF694485 = NC_016576) (Briddon 2001; Wyant et al. 2015; Brown et al. 2015; Zerbini et al. 2017).

West African Asystasia virus 2

(WAAV2)

Synonyms Asystasia begomovirus 2 Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution WAAV2-infected plants of Asystasia gangetica were reported from West Africa (Wyant et al. 2015).

Asystasia gangetica (Chinese violet)

217

Symptoms and host(s) The virus-infected Chinese violet plants exhibit vein yellowing, mosaic, and leaf curling symptoms. Transmission WAAV2 (possibly in combination with WAAV1) has been shown to be transmitted by a specific Bemisia tabaci “type” which transmitted other begomoviruses poorly (Bedford et al. 1992). In common with all other begomoviruses, WAAV2 is likely transmitted by the whitefly in a circulative, nonpropagative manner. Experimentally WAAV2 has been shown to infect Nicotiana benthamiana (exhibiting stunting) and Datura stramonium (exhibiting leaf curling and stunting) by biolistic inoculation of viral DNA. Virion properties and genome The structure of the virions of WAAV2 has not been investigated. In common with all geminiviruses, the virions of WAAV1 are likely geminate (twinned icosahedra). The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2744 nt (JF694486 = NC_038476) (Briddon 2001; Brown et al. 2015; Wyant et al. 2015; Zerbini et al. 2017) including six genes. The expression and function of these genes have not been investigated for WAAV2.

West African Asystasia virus 3 Taxonomic position Genus: Begomovirus

(WAAV3)

Family: Geminiviridae

Geographical distribution WAAV3 infection in plants of Asystasia gangetica was reported from Benin (Leke et al. unpublished – KT444609). Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2728 nt (KT444609) (Brown et al. 2015; Zerbini et al. 2017).

References Bedford ID, Briddon RW, Markham PG, Brown JK, Rossell RK (1992) Bemisia tabaci – biotype characterisation and threat of this whitefly species to agriculture. Proceedings of the Brighton crop protection conference – pests and diseases 1992. pp 1235–1240 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 De Bruyn A, Harimala M, Hoareau M, Ranomenjanahary S, Reynaud B, Lefeuvre P, Lett J-M (2015) Asystasia mosaic Madagascar virus: a novel bipartite begomovirus infecting the weed Asystasia gangetica in Madagascar. Arch Virol 160:1589–1591

A

218

Atractylodes macrocephala (Bai zhu)

Wyant P, Strohmeier S, Fischer A, Schafer B, Briddon RW, Krenz B, Jeske H (2015) Light-dependent segregation of begomoviruses in Asystasia gangetica leaves. Virus Res 195:225–235 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Atractylodes macrocephala (Bai zhu) Family: Asteraceae

Medicinal plant

Atractylodes mild mottle virus Taxonomic position Genus: Caulimovirus

(AMMV)

Family: Caulimoviridae

Geographical distribution AMMV infection in plants of Atractylodes macrocephala was reported from Korea (Lim et al. 2015). Symptoms and host(s) The virus-infected bai zhu plants exhibit mild mottling symptoms. Transmission The virus is transmitted by aphid vectors in a semi-persistent manner. The virus is mechanically saptransmissible. Virion properties and genome The virions are isometric, non-enveloped, and 45–50 nm in diameter with no obvious surface structure. The genome is a monopartite, circular, double-stranded DNA of 8105 bp (KR080327) and contains 6 open reading frames (Hohn 2011).

Atractylodes mottle virus Taxonomic position Genus: Carlavirus

(AtrMV)

Family: Betaflexiviridae

Geographical distribution AtrMV infection in plants of Atractylodes macrocephala was reported from South Korea (Zhao et al. 2015). Symptoms and host(s) The virus-infected bai zhu plants exhibit mottling symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner and also by mechanical sap inoculation.

Atriplex micrantha (Twoscale saltbush)

219

Virion properties and genome The virions are flexuous filaments of about 640 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear positive-sense single-stranded RNA of 8881 nt (KR349343 = NC_038966), excluding the poly(A) tail, and consists of six open reading frames (Zhao et al. 2015).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Atractylodes macrocephala was reported from China (Niu et al. 2014). The virus-infected bai zhu plants exhibit mosaic mottling and dwarfing symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

References Hohn T (2011) Caulimovirus. Caulimoviridae. In: The Springer index of Viruses. Springer, New York, pp 271–277. https://doi.org/10.1007/978-0-387-95919-1_41 Lim S, Igori D, Zhao F, Yoo RH, An TJ, Lim HS, Lee SH, Moon JS (2015) Complete genome sequence of a tentative new caulimovirus from the medicinal plant Atractylodes macrocephala. Arch Virol 160:3127–3131 Niu Y-B, Shi X-L, Zhao H-Q, Zhang X-M, Zhao B-J (2014) Molecular identification and partial sequence analysis of Cucumber mosaic virus isolate from Atractylodes macrocephala Koidz. Acta Phytopathol Sin 44(4):357–362 Zhao F, Igori D, Lim S, Yoo RH, Lee SH, Moon JS (2015) Nucleotide sequence and genome organization of Atractylodes mottle virus, a new member of the genus Carlavirus. Arch Virol 160(11):2895–2898

Atriplex micrantha (Twoscale saltbush) Synonyms Atriplex heterosperma Family: Amaranthaceae

Weed host

Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

IYSV infection in plants of Atriplex micrantha was reported from Utah (USA) (Evans et al. 2009). The virus-infected two-scale saltbush plants exhibit symptoms of spotting and chlorosis, and necrosis on the leaves. The virus is transmitted by a thrips vector, Thrips tabaci, in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of IYSV, refer to Iris spp. or Allium cepa.

A

220

Atropa belladonna (Belladonna)

Sowbane mosaic virus

(SoMV)

Taxonomic position Genus: Sobemovirus

Family: Solemoviridae

SoMV infection in plants of Atriplex suberecta was reported from Australia (Guy 1982). The virus is mechanically sap-transmissible. For more details of SoMV, refer to Chenopodium spp.

References Evans CK, Bag S, Frank E, Reeve J, Ransom C, Drost D, Pappu HR (2009) Natural infection of Iris yellow spot virus in twoscale saltbush (Atriplex micrantha) growing in Utah. Plant Dis 93:430 Guy P (1982) A disease of Atriplex caused by Sowbane mosaic virus. Aust Plant Pathol 11:51–52

Atropa belladonna (Belladonna) Family: Solanaceae

Medicinal

Atropa belladonna virus

(AtBV)

Taxonomic position AtBV is a tentative member of the family Rhabdoviridae. Geographical distribution AtBV infection in plants of Atropa belladonna was first reported from Germany by Lesemann (1972). Symptoms and host(s) The symptoms of virus-infected belladonna plants which are unknown. Virion properties and genome The virions are rhabdo- or bullet-shaped, with a clear modal length of 110 nm and 55 nm wide. The genome is not segmented and contains a single molecule of linear single-stranded RNA.

Belladonna mottle virus Taxonomic position Genus: Tymovirus

(BeMV)

Family: Tymoviridae

Geographical distribution BeMV was first reported in plants of Atropa belladonna from Germany by Bode and Marcus (1959). The virus spreads in Bulgaria and Germany (Paul et al. 1968; Franova 2000).

Aubrieta spp.

221

Symptoms and host(s) The virus-infected belladonna plants exhibit mottling and distortion symptoms (Guy et al. 1984).

A Transmission The virus is transmitted by a beetle vector, Epitrix atropae, in a semi-persistent manner and is not transmitted by aphids. The virus is transmissible by mechanical sap-inoculation but not by contact between plants or by seed. Virion properties and genome The virions are non-enveloped, isometric capsid with T = 3 icosahedral symmetry, about 30 nm in diameter. The genome consists of a single molecule of positive-sense ssRNA. The 30 terminus has a tRNA-like structure. A partial genome sequence of 615 nt is available (X54529) (Ding et al. 1990; Martelli et al. 2002).

References Bode O, Marcus O (1959) Untersuchungen uber eine Virose von Atropa belladonna L., vol I. Verh. IV. intern. Pflanzenschutz-Kongr, Hamburg, 1957, pp 375–376 Ding S, Howe J, Keese P, Mackenzie A, Skotnicki M, Gibbs A (1990) Nucleotide sequence of the virion protein gene of Belladonna mottle tymovirus. Nucleic Acids Res 18:6138 Franova J (2000) Electron microscopy of two viruses of deadly nightshade (Atropa belladonna L.). Acta Virol 44:47–51 Guy PL, Dale JL, Adena MA, Gibbs A (1984) A taxonomic study of the host ranges of tymoviruses. Plant Pathol 33:337–346 Lesemann D (1972) Nachweis eines bazilliformen virus in Atropa belladonna. J Phytopathol 73:83–86 Martelli GP, Sabanadzovic S, Abou-Ghanem Sabanadzovic N, Edwards MC, Dreher T (2002) The family Tymoviridae. Arch Virol 147(9):1837–1846 Paul HL, Bode O, Jankulowa M, Brandes J (1968) Untersuchungen uber ein neues isometrisches virus aus Atropa belladonna L., vol 61. I. Symptomotologie, Reinigung, Morphologie, physikalische und chemische Eigenschaften Phytopathol Z, pp 342–361

Aubrieta spp. Family: Brassicaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Aubrieta spp. was reported from New Zealand (Fletcher 1987). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

222

Aucuba japonica (Spotted laurel)

References Fletcher JD (1987) New plant disease records in New Zealand: additional hosts of Alfalfa mosaic virus and Cucumber mosaic virus. N Z J Agric Res 30:505–506

Aucuba japonica (Spotted laurel) Family: Garryaceae

Ornamental

Aucuba bacilliform virus

(AuBV)

Synonyms Aucuba ringspot virus Taxonomic position AuBV is a tentative member of the genus Badnavirus and family Caulimoviridae Geographical distribution AuBV was first reported in plants of Aucuba japonica by Kusunoki (1980) from Japan. The virus spreads in the UK and New Zealand (Pearson et al. 2006; Veerakone et al. 2015). Symptoms and host(s) The virus-infected spotted laurel plants exhibit yellow ring-spots and mosaic symptoms. Transmission The virus is not transmissible by mechanical inoculation, is transmissible by grafting, is not transmissible by contact between plants, is not transmitted by seed, and is not transmitted by pollen. Virion properties and genome The virions are bacilliform and non-enveloped, c. 130 nm in length and 30 nm wide. The genome is double-stranded DNA.

References Kusunoki M (1980) A fost raportat prima data la Aucuba japonica in Japonia de. Ann Phytopath Soc Jpn 46:414 Pearson MN, Clover GRG, Guy PL, Fletcher JD, Beever RG (2006) A review of the plant virus, viroid, and mollicute records for New Zealand. Aust Plant Pathol 35:217–252 Veerakone S, Tang JZ, Ward LI, Liefting LW, Perez-Egusquiza Z, Lebas BSM, Delmiglio C, Fletcher JD, Guy PL (2015) A review of the plant virus, viroid, liberibacter and phytoplasma records for New Zealand. Australas Plant Pathol 44:463–514

Avena sativa (Oat)

223

Avena fatua (Common wild oat) Family: Poaceae

Barley stripe mosaic virus Taxonomic position Genus: Hordeivirus

A

Weed host

(BSMV)

Family: Virgaviridae

BSMV infection in plants of Avena fatua was reported from Manitoba (Chiko 1975). The virus-infected common wild oat plants exhibit pale green spindle-shaped stripe symptoms. No natural vectors of the virus are known. The virus is transmitted by mechanical sap-inoculation. The virus is seed-transmitted in Avena fatua (Chiko 1975). For more details of BSMV, refer to Hordeum vulgare.

References Chiko AW (1975) Natural occurrence of Barley stripe mosaic virus in wild oats (Avena fatua). Can J Bot 53:417–420

Avena sativa (Oat) Family: Poaceae

Cereal

Barley yellow dwarf virus

(BYDV)

Synonyms Oat red leaf virus Taxonomic position BYDV is a tentative member of the genus Luteovirus and family Luteoviridae BYDV infection in plants of Avena sativa was reported from Poland, Sweden, Africa, Asia, Greece, Australia, Europe, New Zealand, the Czech Republic, North America, and South America (Panayotou 1980; Hoppe et al. 1983; Azzam and D’Arcy 1989; Valverde and Harrison 1989; Ryden 1990). The virus-infected oat plants initially show faint yellowish-green blotches which near the leaf tip, on the margin, or on the leaf blade. Bright yellow discoloration occurs on older but not the youngest leaves except in late infections when only flag leaves show symptoms. Because of the varied color changes during disease development, this disease is also known as “red leaf” (Slykhuis 1967; Goulart et al. 1989). The virus is transmitted by a number of aphid vectors, viz., Rhopalosiphum maidis, R. padi, Schizaphis graminum, Myzus persicae, Macrosiphum avenae, Aulacorthum circumflexum, and Siphonophora circumflexa in a persistent manner. For more details of BYDV, refer to Hordeum vulgare.

224

Avena sativa (Oat)

Barley yellow dwarf virus MAV Taxonomic position Genus: Luteovirus

(BYDV-MAV)

Family: Luteoviridae

BYDV-MAV infection in plants of Avena sativa was reported from the Czech Republic (Kundu 2009). The virus-infected oat plants exhibit yellowing or reddening symptoms. The virus is transmitted by an aphid vector, Rhopalosiphum padi, in a circulative non-propagative manner. The virus is not mechanically sap-transmissible. For more details of BYDV-MAV, refer to Hordeum vulgare.

Barley yellow dwarf virus PAV Taxonomic position Genus: Luteovirus

(BYDV-PAV)

Family: Luteoviridae

BYDV-PAV infection in plants of Avena sativa was reported from Pennsylvania (Gildow and Frank 1988). The virus-infected oat plants exhibit yellowing or reddening symptoms. The virus is transmitted by aphid vectors, Rhopalosiphum padi and Sitobion avenae, in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. For more details of BYDV-PAV, refer to Hordeum vulgare.

Barley yellow striate mosaic cytorhabdovirus Taxonomic position Genus: Cytorhabdovirus

(BYSMV)

Family: Rhabdoviridae

BYSMV infection in plants of Avena sativa was reported from Turkey (Makkouk et al. 1996). The virus-infected oat plants exhibit symptoms of fine, broken chlorotic striations to complete chlorosis. The virus is transmitted by planthopper vectors in a persistent, propagative manner. The virus is not transmitted by mechanical inoculation or through seed. For more details of BYSMV, refer to Hordeum vulgare.

Cereal chlorotic mottle virus

(CCMoV)

Taxonomic position CCMoV is a tentative member of the genus Nucleorhabdovirus and family Rhabdoviridae CCMoV infection in plants of Avena sativa was reported from Australia, Northern Africa, and Morocco (Lockhart 1986). The virus-infected oat plants exhibit symptoms including chlorotic streaking, variable stunting, and sterility. Generally, severe necrotic and chlorotic streaks occur on leaves. The virus is transmitted by leafhopper vectors, Nesoclutha pallida and Cicadulina bimaculata, in a persistent, propagative manner. The virus is not mechanically sap-transmissible and not transmitted through seed. For more details of CCMoV, refer to Hordeum vulgare.

Avena sativa (Oat)

225

Cereal yellow dwarf virus RPV Taxonomic position Genus: Polerovirus

(CYDV-RPV)

Family: Luteoviridae

CYDV-RPV infection in plants of Avena sativa was reported from Alaska and Latvia (Robertson 2003; Bisnieks et al. 2006). The virus-infected oat plants exhibit bright yellow mosaic symptoms. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. For more details of CYDV-RPV, refer to Hordeum vulgare.

Maize dwarf mosaic virus Taxonomic position Genus: Potyvirus

(MDMV)

Family: Potyviridae

MDMV infection in plants of Avena sativa was reported from the United States (McDaniel and Gordon 1989). The virus-infected oat plants exhibit mild mottling symptoms. The virus is transmitted from maize to oat by several aphid species. The virus is also mechanically sap-transmissible. For more details of MDMV, refer to Zea mays.

Maize streak virus

(MSV)

Synonyms African cereal streak virus Taxonomic position Genus: Mastrevirus

Family: Geminiviridae

MSV infection in plants of Avena sativa was reported from East Africa (Harder and Bakker 1973; Karavina 2014). The virus-infected oat plants exhibits symptoms initially as faint, broken, and chlorotic streaks near leaf bases and extend upward. The broken nature of young streaks is clearly defined. Later, definite alternate yellow and green streaks develop along the entire leaf blades. Eventually, leaves become almost completely chlorotic. New leaves tend to develop a shoestring habit and die, and the virus is limited to the phloem where it induces necrosis. Seed yield is almost completely suppressed. Plants become soft, flaccid, and velvety to the touch. The virus is transmitted by the leafhopper vectors, Cicadulina mbila, C. arachidis, C. bipunctella, C. triangular, C. bimaculata, C. similes, C. lateens, C. ghaurii, and C. parazeae, in a persistent, circulative and non-propagative manner. The virus is not transmitted by mechanical sap-inoculation. For more details of MSV, refer to Zea mays.

Moroccan pepper virus Taxonomic position Genus: Tombusvirus

(MPV)

Family: Tombusviridae

A

226

Avena sativa (Oat)

MPV infection in plants of Avena sativa was reported from Southern Iran (Alemzadeh and Ghorbani 2016). The virus is transmitted by mechanical inoculation to three to nine families. There is no vector is known for this virus. For more details of MPV, refer to Capsicum annuum.

Oat blue dwarf virus

(OBDV)

Taxonomic position Genus: Marafivirus

Family: Tymoviridae

Geographical distribution OBDV infection in plants of Avena sativa was first observed in Minnesota, USA, by Goto and Moore (1952). The virus spreads in Illinois, Canada, Finland, Sweden, Europe, and the USA (Banttari and Moore 1962; Banttari 1981). Symptoms and host(s) The virus-infected plants are dwarfed and uniformly dark blue-green and mature later than healthy plants. The dwarfed condition causes diseased plants to be overlooked. Leaves, especially flag leaves, and stems are stiffer and shorter and stand out at a greater angle from the stem than those of healthy plants. Enations along the leaf veins and also sterility of florets are noticed. Tillers appear in larger number and form above the crown. Severely infected heads produce little or no seed. The virus is known to infect both monocot and dicot plants (Westdal 1968; Timian 1985). Transmission The virus is transmitted by aster leafhopper vectors, Macrosteles fascifrons and M. laevis, in a persistent propagative manner. Leafhoppers acquire the virus within 15 min, and the minimum latent period of OBDV is 5 and 8 days in female and male M. fascifrons, respectively. The rate of transmission gradually increases after the latent period, and then after 30 days of acquisition feeding, transmission diminishes. The virus multiplies in leafhoppers; however it is not transovarially transmitted. The virus is not sap or seed-transmissible (Banttari and Zeyen 1970, 1976; Long and Timian 1971). Virion properties and genome The virions are non-enveloped, isometric capsid with T = 3 icosahedral symmetry, about 30 nm in diameter. The genome is a monopartite, linear, positive-sense, single-stranded RNA of 6509 nt (U87832 = NC_001793). There is no VPg at the 50 -terminus (Banttari 1981; Edwards et al. 1997; Martelli et al. 2002; Hammond et al. 2011; Edwards and Weiland 2014).

Oat chlorotic stunt virus Taxonomic position Genus: Avenavirus

(OCSV)

Family: Tombusviridae

Geographical distribution OCSV infection in plants of Avena sativa was first recorded at the Welsh Plant Breeding Station in Aberystwyth, UK (Catherall 1986). The virus was later detected in Shropshire, England (Boonham et al. 1997).

Avena sativa (Oat)

227

Symptoms and host(s) The virus-infected oat plants exhibit symptoms of severe stunting of plants and yellow streaks on the leaves. The chlorotic streaks subsequently turn necrotic. The emerging leaves of oat are dark green and become twisted with age. The severely affected plants do not produce viable seeds. Oat is the natural host. Barley, maize, and wheat are experimental latent hosts. Transmission The virus-infected oat plants are confined in the field in waterlogged areas. Oat seedlings which were grown in soil from infected sites became infected with the virus. This demonstrated that the virus was soil-borne (Boonham et al. 1997). A specific vector of the virus has not been identified so far, but it is assumed that the virus is transmitted by soil-borne zoosporic fungus, Polymyxa olpidium. The virus is readily transmitted mechanically from oat plant to oat plant. Virion properties and genome The virions are isometric, non-enveloped, and 35 nm in diameter and have T = 3 icosahedral symmetry. The genome is monopartite, linear, positive-sense, single-stranded RNA of 4114 nt with three ORFs (X83964 = NC_003633) (Boonham et al. 1995, 1998; Boonham and Flint 2011).

Oat dwarf virus

(ODV)

Taxonomic position Genus: Mastrevirus

Family: Geminiviridae

Geographical distribution ODV infection in plants of Avena sativa was reported from Germany (Schubert et al. 2007). Symptoms and host(s) ODV-infected oat plants exhibited yellowing at the leaf edges and leaf tips early during infection which progressed to stunting and orange-red leaves (Schubert et al. 2007). The ODV virus sequences from Iran were amplified from leafhoppers (species not identified); thus the host cannot be specifically identified (Kamali et al. 2017). Transmission ODV has been shown to be transmissible oat to oat by the leafhopper Psammotettix alienus (Schubert et al. 2007). The mechanism of transmission was not investigated but is likely to be circulative, and nonpropagative, and in common with all other begomoviruses. Virion properties and genome The structure of the virions of ODV has not been investigated. In common with all geminiviruses, the virions of ODV are likely geminate (twinned icosahedra). The genome of ODV consists of a single component of circular single-stranded DNA of ~2740 nt (AM296025 = NC_010799) (Schubert et al. 2007; Kamali et al. 2017; Zerbini et al. 2017). In common with all mastreviruses, ODV is predicted to encode four gene products, two in each orientation. Based on sequence homology to previously studied mastreviruses, these would code for the coat protein and movement protein in the virion-sense and in the complementary-sense for the replication-associated protein (Rep) and Rep A (Palmer and Rybicki 1998; Schubert et al. 2007; Boulton and Davies 2011; Muhire et al. 2013; Kamali et al. 2017).

A

228

Avena sativa (Oat)

Oat golden stripe virus Taxonomic position Genus: Furovirus

(OGSV)

Family: Virgaviridae

Geographical distribution OGSV infection in plants of Avena sativa was first recorded from England and Wales (Plumb et al. 1977). The virus was also reported from Europe and the USA (MacFarlane and Plumb 1978; Adams et al. 1988). Symptoms and host(s) The virus-infected oat plants show yellow stripes on young leaves, although many leaves remain symptomless. Oat is the only natural host. Wheat and barley are not hosts of this virus. Transmission The virus is soil-borne and transmitted by the fungus, Polymyxa betae. P. betae survives in field soil as cystosori. Cysts give rise to zoospores, which swim through free soil water until they contact a host root and encyst. Encysted zoospores produce a structure called a satchel through which zoosporic cytoplasm enters the host cell and becomes a plasmodium. The virus is transmitted through sap-inoculation to Nicotiana benthamiana, N. clevelandii, N. debneyi, and Chenopodium amaranticolor. Virion properties and genome The virions are non-enveloped hollow rods, which have helical symmetry. Two types of virion are composed of rods of about 140–160 nm and 260–300 nm in length and 20 nm in diameter. The genome consists of two segments of positive single-stranded RNA. RNA-1 is 7111 nt long (AJ132578 = NC_002358) and contains three ORFs: ORF-1 which encodes methyltransferase and helicase, ORF-2 RNA-dependent RNA polymerase, and ORF-3 movement protein. RNA-2 is 3232 nt (AJ132579 = NC_002357) and contains three ORFs: ORF-1 encodes 19 kDa coat protein, the ORF is terminated with an opal (UGA) codon; on suppression of this codon, a large coat protein of ORF-2 may be formed. ORF-3 encodes an 18.7 kDa cysteine-rich protein (Adams et al. 1988, 2017; Diao et al. 1999; Adams 2011b).

Oat mosaic virus

(OMV)

Synonyms Soil-borne oat mosaic virus Taxonomic position Genus: Bymovirus

Family: Potyviridae

Geographical distribution OMV infection in plants of Avena sativa was first recorded in South Carolina, USA, in 1944 (Atkinson 1945). The virus spreads in the USA, France, Ireland, Italy, the UK, and New Zealand (MacFarlane et al. 1968; Catherall and Hayes 1970; Monger et al. 2001; Clover et al. 2002).

Avena sativa (Oat)

229

Symptoms and host(s) The virus causes disease in winter oats. In the field mottling symptoms are common in the early stage of growth; subsequently, the symptoms reduce with the further growth and warmer weather. Towards the end of the season, the disease is characterized by stunting of plants. Severely infected plants produce no grain. The natural host range for this virus is restricted to Avena. Transmission The virus is transmitted by Polymyxa graminis, a soil-borne fungus, in a persistent manner (MacFarlane et al. 1968). The virus is mechanically transmitted with difficulty. The virus is not seed-transmitted. Virion properties and genome The virions are non-enveloped, flexuous filaments of two modal lengths, 250–300 nm and 500–600 nm; both are 13 nm in width. The genome consists of two molecules of linear positive-sense ssRNA. RNA-1 is 7550 nt (AJ306718 = NC_004016) with one large open reading frame encoding a polyprotein of 262.8 kDa. RNA2 is 2284 nt (AJ306719 = NC_004017) and substantially smaller than those of other bymoviruses (Usugi and Saito 1981; Zheng et al. 2002; Lopez et al. 2009; Adams 2011a; Wylie et al. 2017).

Oat necrotic mottle virus Taxonomic position Genus: Tritimovirus

(ONMV)

Family: Potyviridae

Geographical distribution ONMV infection in plants of Avena sativa was first recorded in Manitoba, Canada (Gill 1967). The virus spreads in the USA and Canada (Gill 1976; Stenger and French 2004). Symptoms and host(s) The initial symptoms of virus-infected oat plants appear as chlorotic streaks on emerging leaves. Subsequently, irregular light and dark green mottling develops, which contains necrotic lines and irregularly shaped necrotic areas. Similar symptoms also develop in leaf sheaths. Warm weather favors more necrotic symptoms. The natural hosts of this virus are only plants of the Gramineae family. Susceptible hosts are oat cultivars and both wild and cultivated grasses in the Bromus, Lolium, and Poa genera. Transmission The virus is transmitted by mite vectors (possibly). The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, 690–700 nm long and 12–15 nm in diameter. The genome is a monopartite, linear, positive-sense ssRNA of 9436 nt (AY377938 = NC_005136), with a 30 -poly(A) terminus (Gill 1976; Stenger and French 2004; Rabenstein et al. 2002; Lopez-Moya et al. 2009; French et al. 2011; Wylie et al. 2017).

A

230

Avena sativa (Oat)

Oat sterile dwarf virus Taxonomic position Genus: Fijivirus

(OSDV)

Family: Reoviridae

Geographical distribution OSDV infection in plants of Avena sativa was first recorded in Ceskomoravska vrchovina (Czech Republic) and the Northwest district of Slovakia (Prusa et al. 1959; Brcak et al. 1972). The virus is known to be distributed in European countries: Britain, Czechoslovakia, Germany, Sweden, Finland, Norway, and Poland (Lindsten 1961; Catherall 1970; Boccardo and Milne 1980). Symptoms and host(s) The virus induces grasslike appearance in affected plants. Initially the leaves become dark green; subsequently enations may occur on the under surface of the leaves. The affected plants produce excessive stunted tillers. Flowers are mostly sterile resulting in loss of grains (Brcak et al. 1972). Other than oat, barley, maize, rye, and wheat are important natural hosts. Other hosts include: Arrhenatherum elatius, Cynosurus cristatus, Lolium spp., Phalaris canariensis, and Poa annua. L. perenne is the perennial source of infection. Oat and L. multiflorum are good assay hosts. No local lesion host is known for this virus. Transmission The virus is transmitted mainly by the planthopper vector, Javesella pellucida, and also by J. dubia, J. discolor, J. obscurella, and Dicranotropis hamata in a circulative and propagative manner (Boccardo and Milne 1980). The shortest acquisition is 30–60 min., the latent period is 3–4 weeks, and the vectors remain viruliferous throughout life (Vacke 1966). No transovarial or seed transmission of virus has been identified. Virion properties and genome The virions are non-enveloped, icosahedral with a double capsid structure, and about 65–70 nm in diameter. Pentameric turrets sit on the outside of the innermost capsid. The outer capsid has a T = 13 icosahedral symmetry, the inner capsid a T = 2 icosahedral symmetry. The genome is a linear doublestranded RNA which contains 10 segments coding for 12 proteins. The sequences of four genome segments are available: Seg7 is 1944 bp (AB011024 = NC_038655), Seg8 is 1874 bp (AB011025 = NC_038656), Seg9 is 1893 bp (AB011026 = NC_038653), and Seg10 is 1761 bp (AB011027 = NC_038654) (Luisoni et al. 1979; Boccardo and Milne 1980; Francki and Boccardo 1983; Isogai et al. 1998; Harding and Dale 2011).

Oat striate mosaic virus

(OSMV)

Taxonomic position OSMV is a tentative member of the genus Cytorhabdovirus and family Rhabdoviridae Geographical distribution OSMV infection in plants of Avena sativa was first reported in 1975 in Illinois, USA (Jedlinski 1976). The virus spreads in Canada and the Central USA.

Avena sativa (Oat)

231

Symptoms and host(s) The virus-infected plants exhibit striated mosaic and necrosis symptoms on the leaves, and the plants are stunted due to shortened internodes. Transmission The virus is transmitted by the leafhopper vector, Graminella nigrifrons, in a persistent, propagative manner. The vectors acquire the virus during feeding for 1–2 days on infected oats, and an incubation period of 24 days is required before transmission. The vectors acquire the virus both as nymphs and adults. No transovarial transmission is reported. Virion properties and genome The virions are bacilliform or bullet-shaped and measure 400  100 nm (Jedlinski 1976).

Pelargonium leaf curl virus Taxonomic position Genus: Tombusvirus

(PLCV)

Family: Tombusviridae

PLCV infection in plants of Avena sativa was reported from Southern Iran (Alemzadeh and Ghorbani 2016). The virus-infected oat plants exhibit leaf curling symptoms. The virus is mechanically saptransmissible. There is no known vector for this virus. For more details of PLCV, refer to Pelargonium spp.

References Adams M (2011a) Bymovirus. Potyviridae. In: The Springer index of Viruses. Springer, New York, pp 1411–1416. https://doi.org/10.1007/978-0-387-95919-1_232 Adams MJ (2011b) Furovirus. In: Tidona C, Darai G (eds) The Springer index of viruses. Springer, New York. https://doi. org/10.1007/978-0-387-95919-1 Adams MJ, Jones P, Swaby AG (1988) Purification and some properties of Oat golden stripe virus. Ann Appl Biol 112:285–290 Adams MJ, Adkins S, Bragard C, Gilmer D, Li D, MacFarlane SA, Wong S-M, Melcher U, Ratti C, Ryu KH, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Virgaviridae. J Gen Virol 98:1999–2000 Alemzadeh E, Ghorbani A (2016) Occurrence of Pelargonium leaf curl virus and Moroccan pepper virus on natural hosts. Aust Plant Dis Notes 11:8 Atkinson R (1945) A new mosaic chlorosis of oats in the Carolinas. Plant Dis Reptr 29:86–89 Azzam OI, D’Arcy CJ (1989) Survey of spring oats for Barley yellow dwarf viruses in Illinois. Plant Dis 73:610 Banttari EE (1981) Serological assays for Oat blue dwarf virus. Phytopathology 71:1242–1244 Banttari EE, Moore MB (1962) Virus cause of blue dwarf of oats and its transmission to barley and flax. Phytopathology 52:897–902 Banttari EE, Zeyen RJ (1970) Transmission of Oat blue dwarf virus by the aster leafhopper following natural acquisition or inoculation. Phytopathology 60:399–402 Banttari EE, Zeyen RJ (1976) Multiplication of Oat blue dwarf virus in the aster leafhopper. Phytopathology 66:896–900 Bisnieks M, Kvarnheden A, Turka I, Sigvald R (2006) Occurrence of Barley yellow dwarf virus and Cereal yellow dwarf virus in pasture grasses and spring cereals in Latvia. Acta Agric Scand Sect B-Soil Plant Sci 56:171–178 Boccardo G, Milne RG (1980) Oat sterile dwarf virus. CMI/AAB Description of plant viruses, No. 217 Boonham N, Flint L (2011) Avenavirus. Tombusviridae. In: The Springer index of Viruses. Springer, New York, pp 1881–1884. https://doi.org/10.1007/978-0-387-95919-1_308 Boonham N, Henry CM, Wood KR (1995) The nucleotide sequence and proposed genome organization of Oat chlorotic stunt virus, a new soil-borne virus of cereals. J Gen Virol 76:2025–2034

A

232

Avena sativa (Oat)

Boonham N, Harju V, Wood KR, Henry CM (1997) Infection of oats and other cereals by Oat chlorotic stunt virus in the field and laboratory. Plant Pathol 46:795–799 Boonham NM, Henry CM, Wood KR (1998) The characterization of a subgenomic RNA and in vitro translation products of Oat chlorotic stunt virus. Virus Genes 16:141–145 Boulton MI, Davies JW (2011) Mastrevirus. Geminiviridae. In: The Springer index of Viruses. Springer, New York, pp 597–603. https://doi.org/10.1007/978-0-387-95919-1_83 Brcak J, Kralik O, Vacke J (1972) Virus origin of the oat sterile dwarf disease. Biol Plant (Praha) 14(4):302–304 Catherall PL (1970) Oat sterile dwarf virus. Plant Pathol 19:75–78 Catherall PL (1986) Oat virology. Welsh Plant Breeding Station Annual Report. p 132 Catherall PL, Hayes JD (1970) Oat mosaic virus. Plant Pathol 19:78–81 Clover GRG, Ratti C, Rubies-Autonell C, Henry CM (2002) Detection of European isolates of Oat mosaic virus. Eur J Plant Pathol 108:87–91 Diao A, Chen J, Gitton F, Antoniw JF, Mullins J, Hall AM, Adams MJ (1999) Sequences of European wheat mosaic virus and Oat golden stripe virus and genome analysis of the genus furovirus. Virology 261:331–339 Edwards MC, Weiland JJ (2014) Coat protein expression strategy of Oat blue dwarf virus. Virology 450–451:290–296 Edwards MC, Zhang Z, Weiland JJ (1997) Oat blue dwarf marafivirus resembles the tymoviruses in sequence, genome organization, and expression strategy. Virology 232:217–229 Francki RIB, Boccardo G (1983) The plant Reoviridae. In: Joklik WK (ed) The Reoviridae. Plenum Press, New York, pp 505–563 French RC, Stenger DC, Tatineni S (2011) Tritimovirus. Potyviridae. In: The Springer index of Viruses. Springer, New York, pp 1445–1449. https://doi.org/10.1007/978-0-387-95919-1_237 Gildow FE, Frank JA (1988) Barley yellow dwarf virus in Pennsylvania: effect of the PAV isolate on yield components of Noble spring oats. Plant Dis 72:254–256 Gill CC (1967) Oat necrotic mottle, a new virus disease in Manitoba. Phytopathology 57:302–307 Gill CC (1976) Serological properties of Oat necrotic mottle virus. Phytopathology 66:415–418 Goto S, Moore MB (1952) Some oat diseases in Minnesota 1951. Plant Dis Reptr 38:695 Goulart LR, Ohm HW, Foster JE (1989) Barley yellow dwarf symptom severity in oat affected by plant growth stage at infection and plot type. Crop Sci 29:1412–1416 Hammond RW, Edwards MC, Ramirez P (2011) Marafivirus. Tymoviridae. In: The Springer index of Viruses. Springer, New York, pp 1947–1952. https://doi.org/10.1007/978-0-387-95919-1_319 Harder DE, Bakker W (1973) African cereal streak, a new disease of cereals in East Africa. Phytopathology 63:1407–1411 Harding RM, Dale JL (2011) Fijivirus. Reoviridae. In: The Springer index of Viruses. Springer, New York, pp 1589–1593. https://doi.org/10.1007/978-0-387-95919-1_260 Hoppe W, Ruszkiewicz M, Zielinska L (1983) Occurrence of Barley yellow dwarf virus on oat cultivars in Poland. Zesz Probl Postepow Nauk Rol 291:119–129 Isogai M, Uyeda I, Lindsten K (1998) Taxonomic characteristics of fijiviruses based on nucleotide sequences of the Oat sterile dwarf virus genome. J Gen Virol 79:1479–1485 Jedlinski H (1976) Oat striate mosaic virus, a new virus disease in Illinois spread by the leafhopper, Graminella nigrifrons (Forbes). Proc Am Phytopathol Soc 3:208 Kamali M, Heydarnejad J, Pouramini N, Masumi H, Farkas K, Kraberger S, Varsani A (2017) Genome sequences of beet curly top Iran virus, oat dwarf virus, turnip curly top virus, and wheat dwarf virus identified in leafhoppers. Genome Announc 5(8):e01674–e01616 Karavina C (2014) Maize streak virus: a review of pathogen occurrence, biology and management options for smallholder farmers. Afr J Agric Res 9:2736–2742 Kundu JK (2009) First report of Barley yellow dwarf virus-MAV in oat, wheat, and barley grown in the Czech Republic. Plant Dis 93:964 Lindsten K (1961) Studies on virus diseases of cereals in Sweden I. On the etiology of a serious disease of oats (the “Bollnas disease”). Ann Royal Agric Coll. Sweden 27:137–197 Lockhart BEL (1986) Occurrence of Cereal chlorotic mottle virus in northern Africa. Plant Dis 70:912–915 Long DL, Timian RG (1971) Acquisition through artificial membranes and transmission of Oat blue dwarf virus by Macrosteles fascifrons. Phytopathology 61:1230–1232 Lopez-Moya JJ, Valli A, Garcia JA (2009) Potyviridae. In: Iacobellis NS, Collmer A, Hutcheson SW, Mansfield JW, Morris CE, Murillo J, Schaad D NW, Stead E, Surico G, Ullrich M (eds.). Encyclopaedia of Life Sciences (ELS). Kluwer Academic, Dordrecht, The Netherlands. https://doi.org/10.1002/9780470015902.a0000755.pub2 Luisoni E, Boccardo G, Milne RG, Conti M (1979) Purification, serology and nucleic acid of Oat sterile dwarf virus subviral particles. J Gen Virol 45:651–658 MacFarlane I, Plumb RT (1978) A new soil-borne virus of winter oats (Oat golden stripe virus – OGSV). 3rd Int Cong Pl Pathol., Munchen, Aug 16–18 Macfarlane I, Jenkins JEE, Melville SC (1968) A soil-borne virus of winter oats. Plant Pathol 17:167–170

Axonopus compressus (Carpet-grass)

233

Makkouk KM, Bertschinger L, Conti M, Bolay N, Dusunceli F (1996) Barley yellow striate mosaic rhabdovirus naturally infects cereal crops in the Anatolian plateau of Turkey. J Phytopathol 144:413–415 Martelli GP, Sabanadzovic S, Abou-Ghanem Sabanadzovic N, Edwards MC, Dreher T (2002) The family Tymoviridae. Arch Virol 147(9):1837–1846 McDaniel LL, Gordon DT (1989) Characterization of the oat-infecting strain of Maize dwarf mosaic virus. Phytopathology 79:113–120 Monger WA, Clover GRG, Foster GD (2001) Molecular identification of Oat mosaic virus as a Bymovirus. Eur J Plant Pathol 107:661–666 Muhire B, Martin DP, Brown JK, Navas-Castillo J, Moriones E, Zerbini FM, Rivera-Bustamante R, Malathi VG, Briddon RW, Varsani A (2013) A genome-wide pairwise-identity-based proposal for the classification of viruses in the genus Mastrevirus (family Geminiviridae). Arch Virol 158(6):1411–1424 Palmer KE, Rybicki EP (1998) The molecular biology of mastreviruses. Adv Virus Res 50:183–234 Panayotou PC (1980) Occurrence of Barley yellow dwarf virus in Greece. Phytopathol Mediterr 19:143–144 Plumb RT, Catherall PL, Chamberlain JA, Macfarlane I (1977) A new virus of oats in England and Wales. Ann Phytopathol 9:365–370 Prusa V, Jermoljev E, Vacke J (1959) Oat sterile-dwarf virus disease. Biol Plant 1:223–234 Rabenstein F, Seifers DL, Schubert J, French R, Stenger DC (2002) Phylogenetic relationships, strain diversity and biogeography of tritimoviruses. J Gen Virol 83:895–906 Robertson NL (2003) Luteovirus and Polerovirus found in small grains for the first time in the Matanuska-Susitna region of Alaska. Plant Dis 87:446 Ryden K (1990) Chemical control of barley yellow dwarf in spring oats. In: Burnett PA (ed) 1990 – world perspectives on barley yellow dwarf. CIMMYT, Mexico, pp 468–470 Schubert J, Habekuss A, Kazmaier K, Jeske H (2007) Surveying cereal-infecting geminiviruses in Germany – diagnostics and direct sequencing using rolling circle amplification. Virus Res 127(1):61–70 Slykhuis J (1967) Virus diseases of cereals. Rev Appl Mycol 46:401–429 Stenger DC, French R (2004) Complete nucleotide sequence of Oat necrotic mottle virus: a distinct Tritimovirus species (family Potyviridae) most closely related to Wheat streak mosaic virus. Arch Virol 149:633–640 Timian RG (1985) Oat blue dwarf virus in its plant host and insect vectors. Plant Dis 69:706–708 Usugi T, Saito Y (1981) Purification and some properties of Oat mosaic virus. Ann Phytopathol Soc Jpn 47:581–585 Vacke J (1966) Study of transovarial passage of the Oat sterile-dwarf virus. Biol Plant 8:127–130 Valverde RA, Harrison SA (1989) Barley yellow dwarf viruses infecting oats and wheat in Louisiana. Plant Dis 73:938 Westdal PH (1968) Host range studies of Oat blue dwarf virus. Can J Bot 46:1431–1435 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zheng T, Chen J, Chen JP, Adams MJ (2002) The complete sequence of Oat mosaic virus and evidence for deletion and duplication in RNA2. Arch Virol 147:635–642

Axonopus compressus (Carpet-grass) Family: Poaceae

Fodder crop

Axonopus compressus streak virus Taxonomic position Genus: Mastrevirus

(ACSV)

Family: Geminiviridae

Geographical distribution ACSV infection in plants of Axonopus compressus was reported from Nigeria (Oluwafemi et al. 2014).

A

234

Axonopus compressus (Carpet-grass)

Symptoms and host(s) The virus-infected carpet grass plants exhibit streaking symptoms on the leaves. Transmission The virus is transmitted by leafhopper vectors in a persistent, circulative, and non-propagative manner. The virus is not transmitted by mechanical inoculation. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is a monopartite circular single-stranded DNA of 2858 nt (KJ437671 = NC_023864) (Palmer and Rybicki 1998; Boulton and Davies 2011; Muhire et al. 2013; Oluwafemi et al. 2014; Zerbini et al. 2017).

References Boulton MI, Davies JW (2011) Mastrevirus. Geminiviridae. In: The Springer index of Viruses. Springer, New York, pp 597–603. https://doi.org/10.1007/978-0-387-95919-1_83 Muhire B, Martin DP, Brown JK, Navas-Castillo J, Moriones E, Zerbini FM, Rivera-Bustamante R, Malathi VG, Briddon RW, Varsani A (2013) A genome-wide pairwise-identity-based proposal for the classification of viruses in the genus Mastrevirus (family Geminiviridae). Arch Virol 158(6):1411–1424 Oluwafemi S, Kraberger S, Shepherd DN, Martin DP, Varsani A (2014) A high degree of African streak virus diversity within Nigerian maize fields includes a new mastrevirus from Axonopus compressus. Arch Virol 159:2765–2770 Palmer KE, Rybicki EP (1998) The molecular biology of mastreviruses. Adv Virus Res 50:183–234 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

B

Bambusa spp. (Bamboo) Synonyms Dendrocalamus spp. Family: Poaceae

Bamboo mosaic virus Taxonomic position Genus: Potexvirus

Commercial timber

(BaMV)

Family: Alphaflexiviridae

Geographical distribution BaMV infection in plants of Bambusa spp. was first reported from Brazil (Lin et al. 1979). The virus spreads in Southeast Asia, Brazil, the Philippines, Florida (USA), Australia, China, and Taiwan (Chen 1985; Lin and Chen 1991; Lin et al. 1993, 1995, 2005, 2015, 2016a, b; Elliott and Zettler 1996; Thomas and Dodman 1999). Symptoms and host(s) The virus-infected bamboo plants exhibit symptoms of mosaic, necrotic streaks on culms, and also vascular discoloration (Chen 1985). Transmission No vector is reported for this virus. However Chang et al. (2017) have reported BaMV genomic RNAs were detected inside the bodies of two dipteran insects, Gastrozona fasciventris and Atherigona orientalis, but not in thrips (Scirtothrips dorsalis), and these two dipteran insects are suspected to transmit this disease. The virus is also transmitted through contaminated cutting tools during propagation and harvesting activities. The virus is mechanically sap-transmissible to

© Springer Nature India Private Limited 2019 K. S. Sastry et al., Encyclopedia of Plant Viruses and Viroids, https://doi.org/10.1007/978-81-322-3912-3

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Bambusa spp. (Bamboo)

three to nine families, and experimentally infected plants mostly show mosaic and stunting (Lin et al. 1979). Virion properties and genome The virions are flexuous and rod-shaped with a modal length of 490 nm and 13 nm width. The genome is positive-sense single-stranded RNA of 6366 nt (D26017 = NC_001642) and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type, 18–27 kDa in size (Lin et al. 1977, 1992, 1994, 1995, 2016b, 2017; Yang et al. 1997; Adams et al. 2004; Lee et al. 2011).

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Chang K-C, Chang L-T, Huang Y-W, Lai Y-C, Lee C-W, Liao J-T, Lin N-S, Hsu Y-H, Hu C-C (2017) Transmission of Bamboo mosaic virus in bamboos mediated by insects in the order Diptera. Front Microbiol 8:870 Chen TH (1985) Bamboo mosaic virus associated with a mosaic disease of bamboos in Taiwan. Plant Protect Bull (Taichung) 27:111–116 Elliott MS, Zettler FW (1996) Bamboo mosaic virus detected in ornamental bamboo species in Florida. Proc Fla State Hortic Soc 109:24–25 Lee C-C, Ho Y-N, Hu R-H, Yen Y-T, Wang Z-C, Lee Y-C, Hsu Y-H, Meng M (2011) The interaction between Bamboo mosaic virus replication protein and coat protein is critical for virus movement in plant hosts. J Virol 85:12022–12031 Lin NS, Chen CC (1991) Association of Bamboo mosaic virus (BoMV) and BoMV-specific electron-dense crystalline bodies with chloroplasts. Phytopathology 81:1551–1555 Lin MT, Kitajima EW, Cupertino FP, Costa CL (1977) Partial purification and some properties of Bamboo mosaic virus. Phytopathology 67:1439–1443 Lin MT, Chen CC, Kiang T, Lin WC (1979) Preliminary studies on bamboo mosaic disease in Taiwan. Bull Taiwan For Res Inst. Bulletin No. 317 Lin NS, Lin FZ, Huang TY, Hsu YH (1992) Genome properties of Bamboo mosaic virus. Phytopathology 82:731–734 Lin NS, Chai YJ, Huang TY, Chang TY, Hsu YH (1993) Incidence of Bamboo mosaic Potexvirus in Taiwan. Plant Dis 77:448–450 Lin NS, Lin BY, Lo NW, Hu CC, Chow TY, Hsu YH (1994) Nucleotide sequence of the genomic RNA of Bamboo mosaic potexvirus. J Gen Virol 75:2513–2518 Lin N-S, Lin B-Y, Yeh T-Y, Hsu Y-H (1995) First report of Bamboo mosaic virus and its associated satellite RNA on bamboo in the US. Plant Dis 79:1249 Lin J-W, Hsu Y-H, Tsai C-H (2005) Characterization of the infectivity of Bamboo mosaic virus with its correlation to the in vitro replicase activities in Nicotiana benthamiana. Virus Res 112:77–84 Lin WW, Zhang J, Yang WT, Liu YY, Wan BJ, Xu XL, Wu ZJ (2015) First report of Bamboo mosaic virus infecting bamboo in the mainland of China. Plant Dis 99:1189 Lin W-W, Yang W-T, Zhang J, Liu Y-Y, Wang R-Y, Wu Z-J (2016a) Detection of Bamboo mosaic virus by RT-PCR amplification in Fuzhou and Chengdu. Acta Phytopathol Sin 46(4):469–473 Lin W, Gao F, Yang W, Yu C, Zhang J, Chen L, Wu Z, Hsu Y-H, Xie L (2016b) Molecular characterization and detection of a recombinant isolate of Bamboo mosaic virus from China. Arch Virol 161:1091–1094 Lin W, Yan W, Yang W, Yu C, Chen H, Zhang W, Wu Z, Yang L, Xie L (2017) Characterisation of siRNAs derived from new isolates of Bamboo mosaic virus and their associated satellites in infected ma bamboo (Dendrocalamus latiflorus). Arch Virol 162:505–510 Thomas JE, Dodman RL (1999) The first record of Bamboo mosaic potexvirus from Australia. Aust Plant Pathol 28:337 Yang C-C, Liu J-S, Lin C-P, Lin N-S (1997) Nucleotide sequence and phylogenetic analysis of a Bamboo mosaic potexvirus isolate from common bamboo (Bambusa vulgaris McClure). Bot Bull Acad Sin Taipei 38:77–84

Basella alba (Malabar spinach)

237

Barbilophozia spp. Family: Scapaniaceae

Moss plant

B Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

CGMMV infection in plants of Barbilophozia spp. plants was reported from Argentina Islands, Antarctica (Polischuk et al. 2007). There is no known vector for this virus. The virus is transmitted by mechanical sap-inoculation, and also by contact between plants. For more details of CGMMV, refer to Cucumis sativus.

References Polischuk V, Budzanivska I, Shevchenko T, Oliynik S (2007) Evidence for plant viruses in the region of Argentina Islands, Antarctica. FEMS Microbiol Ecol 59:409–417

Basella alba (Malabar spinach) Synonyms Basella rubra Family: Basellaceae

Leafy vegetable

Basella alba alphaendornavirus 1 Taxonomic position Genus: Alphaendornavirus

(BaEV 1)

Family: Endornaviridae

Geographical distribution BaEV 1 infection in plants of Basella alba was reported from Japan (Okada et al. 2014). Symptoms and host(s) The virus-infected Malabar spinach plants do not exhibit visible symptoms. Transmission The virus is transmitted through seed via both ova and pollen. No horizontal spread has been observed in the field, and no potential vectors have been identified. The virus is not mechanically sap-transmissible.

238

Bauhinia variegata (Mountain ebony, Kachnar)

Virion properties and genome The virus does not have gene for capsid protein; therefore no virion is formed. The genome is linear dsRNA of 14,027 bp (AB844265). The dsRNA genome is encapsulated along with the viral replicase enzyme. A site-specific break (nick) is found in the coding strand about 1–2 kb from the 50 terminus. The genome encodes for one ORF potentially cleaved into several polypeptides.

Basella rugose mosaic virus Taxonomic position Genus: Potyvirus

(BaRMV)

Family: Potyviridae

Geographical distribution BaRMV infection in plants of Basella rubra was reported from China and Taiwan (Huang and Chang 2006; Jhu et al. Unpublished - NC_009741). Symptoms and host(s) The virus-infected Malabar spinach plants exhibit mosaic and rugose symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear positive-sense single-stranded RNA of 9804 nt (DQ821938 = NC_009741) (Revers and Garcia 2015; Wylie et al. 2017).

References Huang C-H, Chang Y-C (2006) Basella rugose mosaic virus, a new potyvirus infecting Basella rubra. Plant Pathol 55:819 Okada R, Kiyota K, Moriyama H, Fukuhara T, Valverde RA (2014) A new endornavirus species infecting Malabar spinach (Basella alba L.). Arch Virol 159:807–809 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354

Bauhinia variegata (Mountain ebony, Kachnar) Family: Fabaceae

Ornamental

Apple stem grooving virus Taxonomic position Genus: Capillovirus

(ASGV)

Family: Betaflexiviridae

Begonia spp.

239

ASGV infection in plants of Bauhinia variegata was reported from India (Bhardwaj et al. 2015). The virus-infected mountain ebony plants exhibit severe virus-like symptoms, which included chlorosis, chlorotic and necrotic spots, mottling, leaf distortion, and grooves on the stem. The virus is mechanically sap-transmissible and by grafting. For more details of ASGV, refer to Malus domestica.

B References Bhardwaj P, Ram R, Zaidi AA, Hallan V (2015) Natural occurrence of Apple stem grooving virus on Bauhinia variegata. Trees 29:1415–1422

Beaucarnea recurvata (Elephant’s foot, Ponytail palm) Family: Dracenaceae

Ornamental

Tomato yellow ring virus

(TYRV)

Taxonomic position TYRV is a tentative member of the genus Orthotospovirus and family Tospoviridae TYRV has been reported to infect Beaucarnea recurvata in Iran (Ghotbi and Shahraeen 2012). The virus is transmitted by thrips vectors in a persistent, propagative manner, and also by mechanical sap-inoculation. For more details of TYRV, refer to Solanum lycopersicum.

References Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses infecting propagated ornamentals in Northern Iran. Int Res J Microbiol 3:373–381

Begonia spp. Family: Begoniaceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

Family: Secoviridae

ArMV infection in plants of Begonia spp. was reported from Belgium and Lithuania (Welvaert and Samyn 1978; Welvaert et al. 1980; Albouy 1995; Samuitiene et al. 2008). The virus-infected plants exhibit symptoms of necrosis, stunting, and death of the plants. This virus is transmitted by nematode

240

Begonia spp.

vectors (Xiphinema spp.) in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Beet ringspot virus

(BRSV)

Taxonomic position Genus: Nepovirus

Family: Secoviridae

BRSV infection in plants of Begonia ricinifolia was reported from Hungary (Kis et al. 2017). The virusinfected begonia plants exhibit chlorotic ringspot and line pattern symptoms. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BRSV, refer to Beta vulgaris.

Begonia flower breaking virus

(BFBV)

Taxonomic position BFBV is a tentative member of the genus Potyvirus and family Potyviridae Geographical distribution BFBV infection in plants of Begonia semperflorens was reported from the Netherlands (Gera and Zeidan 2006; FJ539085, Pham et al. unpublished) and in B. bowerae from China (Wu et al. 2016). Symptoms and host(s) The virus-infected plants exhibit flower breaking symptoms. Transmission The virus is presumed to be transmitted by aphid vectors. Virion properties and genome The virions are flexuous filaments. The genome is a single molecule of positive-sense single-stranded RNA. A partial genome sequence of 1423 nt is available (FJ539085) (Wylie et al. 2017).

Broad bean wilt virus Taxonomic position Genus: Fabavirus

(BBWV)

Family: Secoviridae

BBWV (but without specifying either of the currently recognized species BBWV-1 or BBWV-2) infection in plants of Begonia semperflorens was reported from the USA (Lockhart and Betzold 1982). The virus-infected plants exhibit symptoms of leaf mottling, faint ringspots, and stunting. Color breaking is noticed in pink- and red-flowered varieties. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a number of host plants. For more details of BBWV, refer to Vicia faba.

Begonia spp.

241

Carnation mottle virus

(CarMV)

Taxonomic position Genus: Alphacarmovirus

Family: Tombusviridae

CarMV was detected in plants of Begonia elatior and B. cheimantha in Denmark, showing veinclearing, leaf curl, and flower break; symptoms were most prominent during the winter months and affected saleability (Paludan and Begtrup 1985; Albouy 1995). The virus is transmitted by means not involving a vector. The virus is transmissible by mechanical sap-inoculation, by grafting, and also by contact between plants. For more details of CarMV, refer to Dianthus caryophyllus.

Clover yellow mosaic virus Taxonomic position Genus: Potexvirus

(ClYMV)

Family: Alphaflexiviridae

ClYMV infection in plants of Begonia semperflorens was reported from the USA (Koenig 1985). The virus-infected plants exhibit symptoms of bright mosaic and stunting of the plants. No vectors have been reported for this virus. The virus is transmissible by mechanical sap-inoculation. For more details of ClYMV, refer to Trifolium spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Begonia spp. is worldwide in distribution (Brierley and Travis 1958; Zaidi et al. 2011). The virus-infected plants show leaf symptoms such as mottling, chlorosis, and ringspots. Internodes are shortened and the vigor of the plant is decreased due to stunting. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV is reported on plants of Begonia spp. from Canada, Auckland, Bosnia, and Herzegovina (Southeastern Europe) (Tegel and Lemmetty 1998; Kurokawa et al. 2003; Elliott et al. 2009; Zaidi et al. 2011; Tian et al. 2013; Trkulja et al. 2013). The virus-infected begonia plants exhibit symptoms of necrotic areas on leaves, mosaic patterns, concentric rings, stem necrosis, vein necrosis, stunting, puckering, and “burning” of foliage (German et al. 1992). The virus is transmitted by thrips vectors in a persistentpropagative manner, and also by mechanical sap-inoculation. Cuttings from the virus-infected plants used for vegetative propagation is the primary mode of spread. For more details of INSV, refer to Impatiens spp.

B

242

Begonia spp.

Prunus necrotic ringspot virus Taxonomic position Genus: Ilarvirus

(PNRSV)

Family: Bromoviridae

PNRSV infection on plants of Begonia spp. was reported from India (Verma et al. 2002). The virusinfected plants exhibit chlorotic or necrotic ringspot symptoms. The virus is transmitted by the thrips vectors, is present in/on pollen, and enters into the host through the injuries caused by thrips while feeding. The virus is transmitted through the use of virus-infected vegetative propagative material. The virus is mechanically sap-transmissible, and inoculated Cyamopsis tetragonoloba produced dark-colored local lesions, typical of PNRSV. For more details of PNRSV, refer to Prunus persica.

Tobacco necrosis virus A Taxonomic position Genus: Alphanecrovirus

(TNV-A)

Family: Tombusviridae

TNV-A infection in plants of Begonia spp. was reported from India and Denmark (Kristensen 1967; Paludan and Thomsen 1982; Zaidi et al. 2011). The virus-infected plants exhibit stunting, poor vigor, ringspots, and color variation symptoms. The virus is transmitted by the zoospores of the fungus Olpidium brassicae, and the virus is also mechanically sap-transmissible. For more details of TNV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Begonia spp. was reported from Minnesota (USA) (Lockhart and Betzold 1979). The virus-infected begonia plants exhibited chlorotic or necrotic ringspot symptoms. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Begonia spp. is worldwide in distribution (Barnes and Halliwell 1985; German et al. 1992; Daughtrey 1996; Zaidi et al. 2011). The virus-infected plants exhibited chlorotic or necrotic ringspot symptoms. Plants are stunted and flowers are of poor quality due to virus infection. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Begonia spp.

243

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Begonia spp. was reported from south Florida (Ying and Davis 2000). The virus-infected plants exhibit symptoms of curling, distortion and chlorosis of leaves, stunting of the plant, and a high rate of flower abscission. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not transmissible by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

Zucchini yellow mosaic virus Taxonomic position Genus: Potyvirus

(ZYMV)

Family: Potyviridae

ZYMV infection in plants of Begonia semperflorens was reported from Taichung, Chinese Taipei (Chen and Hong 2008; Hong and Chen 2011). The virus-infected plants exhibited faint ringspots in leaves at the early stage of infection. Symptoms turned chlorotic and coalesced as the disease progressed. The virus is transmitted by a number of aphid species in a non-persistent. The virus is mechanically saptransmissible to a number of hosts, and chlorotic spots were produced on Chenopodium quinoa. Among the susceptible hosts, more than 20 different cucurbit species have been found to be susceptible to this virus. For more details of ZYMV, refer to Cucurbita pepo.

References Albouy J (1995) Begonia. Chapter 41. In: Loebenstein G, Lawson RH, Brunt AA (eds) Virus and virus-like diseases of bulb and flower crops. Wiley, Chichester Barnes LW, Halliwell RS (1985) Tomato spotted wilt virus infecting begonia hybrids in Texas. Plant Dis 69:613 Brierley P, Travis RV (1958) Soil-borne viruses from chrysanthemum and begonia. Plant Dis Reptr 42:1030–1033 Chen YK, Hong YH (2008) First report of begonia chlorotic ringspot caused by Zucchini yellow mosaic virus in Taiwan. Plant Dis 92:1247 Daughtrey ML (1996) Detection and identification of tospoviruses in greenhouses. Acta Hortic 431:90–98 Elliott DR, Lebas BSM, Ochoa-Corona FM, Tang J, Alexander BJR (2009) Investigation of Impatiens necrotic spot virus outbreaks in New Zealand. Aust Plant Pathol 38:490–495 Gera A, Zeidan M (2006) New and emerging virus diseases in ornamental crops. Acta Hortic 722:175–180 German TL, Ullman DE, Moyer JW (1992) Tospoviruses: diagnosis, molecular biology phylogeny and vector relationships. Ann Rev Phytopathol 30:315–348 Hong Y-H, Chen Y-K (2011) Zucchini yellow mosaic virus causes begonia chlorotic ring spot in Chinese Taipei. Acta Hortic 901:173–179 Kis S, Salamon P, Kis V, Szittya G (2017) Molecular characterization of a Beet ringspot nepovirus isolated from Begonia ricinifolia in Hungary. Arch Virol 162:3559–3562 Koenig R (1985) Recently discovered virus or virus-like diseases of ornamentals and their epidemiological significance. Acta Hortic 164:21–32 Kristensen HR (1967) Virus diseases of ornamental plants. Begonia x cheimantha. Plant diseases and pests in Denmark, 83rd annual report: 315–335 Kurokawa E, Kameya-Iwaki M, Okuda M, Kajihara H, Hanada K, Ito S, Tanaka S (2003) Necrotic mottle disease of begonia caused by Impatiens necrotic spot virus (abstract in Japanese). Jpn J Phytopathol 69:67 Lockhart BE, Betzold JA (1979) Begonia yellow spot: a disease caused by Tobacco ringspot virus infection. Plant Dis Reptr 63:1046–1047 Lockhart BEL, Betzold JA (1982) Broad bean wilt virus in begonia in Minnesota. Plant Dis 66:72–73

B

244

Benincasa hispida (Ash gourd/Wax gourd)

Paludan N, Begtrup J (1985) Carnation mottle virus demonstrated in Begonia elatior and Begonia x cheimantha showing vein clearing, leaf curl and flower break. Acta Hortic 164:33–40 Paludan N, Thomsen A (1982) Virus diseases of ornamental plants. Begonia elatior. Plant diseases and pests in Denmark 1981, 98th annual report: 55–56 Samuitiene M, Navalinskiene M, Jackeviciene E (2008) Arabis mosaic virus on ornamental plants. Biologia 54:264–268 Tegel J, Lemmetty A (1998) Palsamin kuoliolaikkuvirus loydetty begoniasta. Puutarha Kauppa 17/98:17 Tian X, Zheng Y, Chintaluri K, Meng B (2013) First report of Impatiens necrotic spot virus on Hiemalis begonia (Begonia  hiemalis) in Canada. Plant Dis 97(2):291 Trkulja V, Mihic Salapura J, Curkovic B, Stankovic I, Bulajic A, Vucurovic A, Krstic B (2013) First report of Impatiens necrotic spot virus on begonia in Bosnia and Herzegovina. Plant Dis 97(7):1004 Verma N, Hallan V, Ram R, Zaidi AA (2002) Detection of Prunus necrotic ringspot virus in begonia by RT–PCR. Plant Pathol 51:800 Welvaert W, Samyn G (1978) The occurrence of Arabis mosaic virus in vegetatively propagated begonias: Begonia tuberhybrida Voss. cv. Multiflora. Meded van de Fac Landbouwwetenschappen Rijksuniversiteit Gent 43:1051–1061 Welvaert W, Samyn G, Van Wymersch E (1980) On the production of virus free Begonia tuberhybrida Voss cv. Multiflora varieties. Acta Hortic 110:253–258 Wu HZ, Gao J, Xu Y, Wu JZ, He SL, Chen HR (2016) First report of Begonia flower breaking virus infecting Begonia bowerae in China. Plant Dis 100:2538 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354 Ying ZT, Davis MJ (2000) Partial characterization and host range of Tomato yellow leaf curl virus in South Florida. Proc Fla State Hortic Soc 113:185–190 Zaidi AA, Hallan V, Raikhy G, Singh AK, Ram R (2011) Viruses of ornamental plants in India – current research status and future prospects. Acta Hortic 901:67–76

Benincasa hispida (Ash gourd/Wax gourd) Family: Cucurbitaceae

Vegetable

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

CGMMV-infected plants of Benincasa hispida were reported from India (Kumar et al. 2017). The virus is transmissible by mechanical sap-inoculation, and by contact between plants. Transmission through contaminated irrigation water and nutrient solutions has also been reported. The virus is also pollen transmissible. For more details of CGMMV, refer to Cucumis sativus.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Benincasa hispida was reported from Western Samoa (Pearson and Liyanage 1997). The virus-infected ash gourd plants exhibit mosaic symptoms. The virus is transmitted

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by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(CABYV)

Family: Luteoviridae

CABYV infection in plants of Benincasa hispida was reported from China and Taiwan (Xiang et al. 2008; Knierim et al. 2010). The virus-infected ash gourd plants exhibit yellowing of the older leaves, and leaves are thickened and brittle. The virus is transmitted by aphids, Myzus persicae, M. euphorbiae, and Aphis gossypii, in a circulative, non-propagative manner (Lecoq et al. 1992) and not transmissible by mechanical sap-inoculation. For more details of CABYV, refer to Cucumis melo.

Melon yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(MYSV)

Family: Tospoviridae

MYSV infection in plants of Benincasa hispida was reported from Japan and China (Okuda et al. 2002). The virus-infected ash gourd plants exhibit leaf and stem necrosis symptoms. The virus is transmitted by the thrips vector, Thrips palmi, in a persistent, propagative manner. For more details of MYSV, refer to Cucumis melo.

Papaya ringspot virus Taxonomic position Genus: Potyvirus

(PRSV)

Family: Potyviridae

PRSV infection in Benincasa hispida plants was reported from Tamilnadu (India) (Nagendran et al 2017). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap inoculation. For more details of PRSV, refer to Carica papaya.

Squash leaf curl China virus Taxonomic position Genus: Begomovirus

(SLCCNV)

Family: Geminiviridae

SLCCNV infection in plants of Benincasa hispida was reported from India and Thailand (Sawangjit, 2009; Mohammed-Riyaz et al. 2013). The virus-infected ash gourd plants exhibited upward leaf curling, crinkling, puckering, yellowing, and reduced lamina. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not mechanically saptransmissible. For more details of SLCCNV, refer to Cucurbita pepo.

B

246

Benincasa hispida (Ash gourd/Wax gourd)

Squash leaf curl Philippines virus Taxonomic position Genus: Begomovirus

(SLCuPV)

Family: Geminiviridae

SLCuPV infection in plants of Benincasa hispida was reported from Taiwan (Liao et al. 2007). The virus-infected ash gourd plants exhibit yellowing, vein enation, rugose mosaic, and leaf curling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not mechanically sap-transmissible. For more details of SLCuPV, refer to Cucurbita pepo.

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Benincasa hispida was reported from India (Roy et al, 2013). The virus-infected ash gourd plants exhibit puckering, yellowing of leaves, and stunting symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner (Tamil Nayagan et al. 2016). The virus is graft transmitted. For more details of ToLCNDV, refer to Solanum lycopersicum.

Tomato leaf curl virus

(ToLCV)

Taxonomic position Genus: Begomovirus

Family: Geminiviridae

ToLCV infection in plants of Benincasa hispida was reported from Thailand (Samretwanich et al. 2000). The virus-infected ash gourd plants exhibit leaf yellowing symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is graft transmitted. For more details of ToLCV, refer to Solanum lycopersicum.

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV was identified infecting plants of Benincasa hispida in Gorakhpur, India (Bhargava and Bhargava 1977). The virus-infected ash gourd plants show mosaic or mottling symptoms. The virus is transmitted by aphid vectors, Myzus persicae and Aphis gossypii, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of WMV, refer to Citrullus lanatus.

Benincasa hispida (Ash gourd/Wax gourd)

247

Watermelon silver mottle orthotospovirus Taxonomic position Genus: Orthotospovirus

(WSMoV)

Family: Tospoviridae

WSMoV infection on plants of Benincasa hispida was reported from Taiwan (Chen et al. 1995; Okuda et al. 2002). The virus-infected ash gourd plants exhibit symptoms of light green to darker green mottle on leaves with waxless surface, leaf crinkling, and tip necrosis. The virus is transmitted by the thrips vector, Thrips palmi, in a persistent-propagative manner, and also by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants. For more details of WSMoV, refer to Citrullus lanatus.

Zucchini yellow mosaic virus Taxonomic position Genus: Potyvirus

(ZYMV)

Family: Potyviridae

ZYMV has been recorded in plants of Benincasa hispida from Japan (Fukumoto et al. 1993). The virusinfected ash gourd plants exhibit yellow mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ZYMV, refer to Cucurbita pepo.

References Bhargava B, Bhargava KS (1977) Cucurbit mosaic viruses in in Gorakhpur. Indian J Agric Sci 47:1–5 Chen CC, Ho HM, Chang TF, Chao CH, Yeh SD (1995) Characterization of a tospovirus-like virus isolated from wax gourd. Plant Protect Bull (Taipei) 37:117–131 Fukumoto F, Terami F, Ishii M (1993) Zucchini yellow mosaic virus isolated from wax gourd (Benincasa hispida Cogn.) and balsam pear (Momordica charantia L.) (in Japanese). Proc Kanto Plant Protect Soc 40:101–103 Knierim D, Deng TC, Tsai WS, Green SK, Kenyon L (2010) Molecular identification of three distinct Polerovirus species and a recombinant Cucurbit aphid-borne yellows virus strain infecting cucurbit crops in Taiwan. Plant Pathol 59:991–1002 Kumar A, Jailani AAK, Roy A, Mandal B (2017) The occurrence, biology and genomic properties of tobamoviruses infecting crop plants in India. In: Mandal B, Rao GP, Baranwal VK, Jain RK (eds) A century of plant virology in India. Springer Publishers, Singapore, pp 429–443 ISBN 978-981-10-5671-0 Lecoq H, Bourdin D, Wipf-Scheibel C, Bon M, Lot H, Lemaire O, Herrbach E (1992) A new yellowing disease of cucurbits caused by a Luteovirus, Cucurbit aphid-borne yellows virus. Plant Pathol 41:749–761 Liao J-Y, Hu C-C, Lin T-K, Chang C-A, Deng T-C (2007) Identification of Squash leaf curl Philippines virus on Benincasa hispida in Taiwan. Plant Pathol Bull 16:11–18 Mohammed-Riyaz SU, Deepan S, Dharanivasan G, Jesse MI, Muthuramalingam R, Kathiravan K (2013) First report on a variant of Squash leaf curl China virus (SLCCNV) infecting Benincasa hispida in India. New Dis Rep 28:20 Nagendran K, Mohankumar S, Aravintharaj R, Balaji CG, Manoranjitham SK, Singh AK, Rai AB, Singh B, Karthikeyan G (2017) The occurrence and distribution of major viruses infecting cucurbits in Tamil Nadu state, India. Crop Protection 99: 10–16 Okuda M, Takeuchi S, Taba S, Kato K, Hanada K (2002) Melon yellow spot virus and Watermelon silver mottle virus: outbreak of cucurbit infecting tospovirus in Japan. Acta Hortic 588:143–148 Pearson MN, Liayanage AS (1997) Records of cucurbit viruses infecting vegetable crops in Western Samoa. Aust Plant Pathol 28:188–191 Roy A, Spoorthi P, Panwar G, Kumar Bag M, Prasad TV, Kumar G, Gangopadhyay KK, Dutta M (2013) Molecular evidence for occurrence of Tomato leaf curl New Delhi virus in ash gourd (Benincasa hispida) germplasm showing a severe yellow stunt disease in India. Indian J Virol 24(1):74–77

B

248

Berberis spp.

Samretwanich K, Chiemsombat P, Kittipakorn K, Ikegami M (2000) Yellow leaf disease of cantaloupe and waxgourd from Thailand caused by Tomato leaf curl virus. Plant Dis 84:200 Sawangjit S (2009) The complete nucleotide sequence of Squash leaf curl China virus-(wax gourd) and its phylogenetic relationship to other geminiviruses. Sci Asia 35:131–136 Tamil Nayagan T, Suganthy M, Renuka Devi P, Malathi VG, Ganapathy N (2016) Vector transmission of tomato leaf curl New Delhi virus in ash gourd and occurrence of Asia II 6 whitefly genotype in India. In: International conference (VIROCON 2016), Bangalore, India, pp 123–124 Xiang HY, Shang QX, Han CG, Li DW, Yu JL (2008) First report on the occurrence of Cucurbit aphid-borne yellows virus on nine cucurbitaceous species in China. Plant Pathol 57:390

Berberis spp. Family: Berberidaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Medicinal

(CMV)

Family: Bromoviridae

CMV infection in plants of Berberis thunbergii was reported from New York (USA) (Wilkinson 1953). The virus-infected plants exhibit leaf mosaic characterized by reddish blotches, varying degrees of foliar distortion and restriction of leaf area. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Berberis spp. was reported from Pennsylvania (USA) (Hausbeck et al. 1992). The virus-infected plants exhibit mottle mosaic, chlorotic spotting, and stunting symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among greenhouse ornamentals in Pennsylvania. Plant Dis 76:795–800 Wilkinson RE (1953) Berberis thunbergii, a host of Cucumber mosaic virus. Phytopathology 43:489

Beta vulgaris (Sugar beet)

249

Berteroa incana (Hoary alyssum) Family: Brassicaceae

Ornamental

B Cherry leaf roll virus Taxonomic position Genus: Nepovirus

(CLRV)

Family: Secoviridae

CLRV infection on plants of Berteroa incana was reported from the USA (Lockhart 1977). The virusinfected plants do not show any symptoms. The virus is transmitted by nematode vectors, Xiphinema coxi, X. diversicaudatum, and X. vuittenezi, in a non-persistent manner, and also by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants. For more details of CLRV, refer to Prunus avium.

References Lockhart BEL (1977) Cherry leaf roll virus in Berteroa incana. Phytopathol News 4:90

Beta vulgaris (Sugar beet) Family: Amaranthaceae

Vegetable

Alfalfa mosaic virus

(AMV)

Taxonomic position Genus: Alfamovirus

Family: Bromoviridae

AMV occurs wherever Beta vulgaris is grown (Shepherd et al. 1965; Farzadfar et al. 2006). The virusinfected sugar beet plants exhibit most characteristic symptoms of the disease consisting of prominent yellow blotches and ringspots. Little, if any, distortion or blistering was present on any of the diseased plants. The virus is transmitted by more than 13 aphid species in a non-persistent manner. The virus is mechanically sap-transmissible and also through pollen to the seed. For more details of AMV, refer to Medicago sativa.

Beet black scorch virus Taxonomic position Genus: Betanecrovirus

(BBSV)

Family: Tombusviridae

250

Beta vulgaris (Sugar beet)

Geographical distribution BBSV infection in plants of Beta vulgaris was reported from Asia, Iran, Europe, China, and the USA (Liu and Xian 1995; Cai et al. 1999; Weiland et al. 2006, 2007; Koenig and Valizadeh 2008; GonzalezVazquez et al. 2009). Symptoms and host(s) The virus-infected sugar beet leaves show diffused chlorotic local lesions and also severe symptoms of black scorch lesions on the leaves. Lateral roots and root hairs of beetroots exhibit rhizomania-like symptoms including necrosis on roots. Host range of the virus for localized and occasionally systemic infection included the Chenopodiaceae and Tetragonia expansa; Nicotiana benthamiana supported symptomless systemic infection by the virus. Transmission The virus spreads by the fungal vector (Olpidium brassicae) in a non-persistent manner (Jiang et al. 1999). The virus is mechanically sap-transmissible, and 13 plant species in 4 families could be locally infected. Among them, Chenopodium amaranticolor, C. quinoa, C. murale, Spinacia oleracea, and Tetragonia expansa show local lesions, while symptomless infections are produced on Lactuca sativa, Physalis floridana, and six Nicotiana species (Cao et al. 2002; Weiland et al. 2007). Virion properties and genome The virions are isometric 28 nm in diameter and have T = 3 icosahedral symmetry. The genome is a monopartite, linear, single-stranded positive-sense RNA of 3644 nt (AF452884 = NC_004452) containing 5 open reading frames (ORFs). The ORF-1 encodes a 23 kDa protein. The ORF-1 contains one readthrough ORF that encodes an 82 kDa protein. ORF-2 to ORF-4 are located in the center of the genome, which encode a 4.2 kDa protein and two 7 kDa proteins. The ORF-5 encodes a 24.5 kDa coat protein (Cao et al. 2002; Yuan et al. 2006).

Beet chlorosis virus Taxonomic position Genus: Polerovirus

(BChV)

Family: Luteoviridae

Geographical distribution BChV infection in plants of Beta vulgaris was reported from Europe and the USA (Hauser et al. 2000, 2002; Stevens et al. 2004; Kozlowska-Makulska et al. 2007). Symptoms and host(s) The virus-infected sugar beet plants exhibit symptoms such as interveinal yellowing, leaf chlorosis, and necrotic lesions (Hauser et al. 2002). Transmission The virus is transmitted by Aphis fabae, Myzus ascalonicus, and M. persicae in a circulative and nonpropagative manner. Vector transmission was successful when a 48-hour acquisition access period and an inoculation access period of 3 days are provided. The virus is not mechanically sap-transmissible.

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251

Virion properties and genome The virions are 26 nm in diameter, are hexagonal in outline, and have no envelope. The genome consists of a single-stranded positive-sense RNA (Stevens et al. 2005). The complete nucleotide sequences of BChV-2a isolate from England and an isolate from California are 5776 nt (AF352024) and 5742 nt (AF352025), respectively. The 30 terminus has neither a poly(A) tract nor a tRNA-like structure. The 50 terminus has a genome-linked protein (VPg) (of about 17 kDa). The genome contains six open reading frames (ORFs) (Hauser et al. 2002).

Beet cryptic virus 1 Taxonomic position Genus: Alphapartitivirus

(BCV-1)

Family: Partitiviridae

Geographical distribution BCV-1 was first reported from the UK on Beta vulgaris and also present in the Australian and Eurasian regions, Japan, and the USA (White and Woods 1978; Xie et al. 1989, 1994). Symptoms and host(s) This virus-infected sugar beet plants do not express any symptoms, and few plants belonging to three families are susceptible. Transmission No insect vector is reported for this virus. The virus is not mechanically sap-transmissible and there is no transmission through contact. The virus is not graft-transmissible. The virus is transmitted by pollen to the seed. 100% seed transmission was recorded in through true seed of beet (Natsuaki et al. 1986). Virion properties and genome The virions are non-enveloped isometric and 30 nm in diameter. The genome consists of two segments of double-stranded RNA. RNA-1 consists of 2008 bp (EU489061 = NC_011556) and RNA-2 1783 bp (EU489062 = NC_011557) (Accotto and Boccardo 1986). The RNA-1 consists of only one major ORF on the plus strand encoding a 72.5 kDa protein that contains sequence motifs characteristic for RNAdependent RNA polymerases (Accotto et al. 1987). The RNA-2 (1783 bp) encodes for the viral coat protein (CP) (Szego et al. 2010; Vainio et al. 2018).

Beet cryptic virus 2 Taxonomic position Genus: Deltapartitivirus

(BCV-2)

Family: Partitiviridae

Geographical distribution BCV-2 was first reported in Beta vulgaris from the UK (Kassanis et al. 1977). The virus has also been found in Australia, the Eurasian region, Japan, and the USA (Xie et al. 1989). Symptoms and host(s) No symptoms are noticed in BCV-2-infected sugar beet plants.

B

252

Beta vulgaris (Sugar beet)

Transmission No insect vector is reported for this virus. The virus is not mechanically sap-transmissible and also no contact transmission. The virus is not transmitted by grafting. The virus is transmitted by pollen to the seed. However, 100% seed transmission was recorded in the true seed of beetroot (Kassanis et al. 1978; White and Woods 1978; Accotto and Boccardo 1986; Natsuaki et al. 1986). Virion properties and genome The virions are isometric, non-enveloped, and 30 nm in diameter. The genome consists of three monocistronic dsRNAs: RNA-1 consists of 1598 bp (HM560702), RNA-2 1575 bp (HM560703), and RNA-3 1522 bp (HM560704). None of the BCV-2 dsRNAs contain a poly(A) tail (Xie et al. 1989; Szego 2009; Vainio et al. 2018).

Beet curly top Iran virus Taxonomic position Genus: Becurtovirus

(BCTIV)

Family: Geminiviridae

Geographical distribution BCTIV infection in plants of Beta vulgaris was reported from Iran (Yazdi et al. 2008; Heydarnejad et al. 2013; Kardani et al. 2013; Soleimani et al. 2013; Varsani et al. 2014a; Anabestani et al. 2016). Symptoms and host(s) The virus-affected sugar beet plants exhibit leaf curling, vein-clearing, and swelling of veins on the lower leaf surface. The virus has been isolated from sugar beet (Beta vulgaris), turnip (Brassica rapa), tomato (Solanum lycopersicum), cowpea (Vigna unguiculata), and bean (Phaseolus vulgaris) (Kardani et al. 2013). Transmission The virus is transmitted by the leafhopper vector, Circulifer haematoceps, in a persistent manner (Taheri et al. 2012). Due to short life cycle and high populations of the leafhopper vector, the disease often spreads very rapidly. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome of the virus is single molecule of circular ssDNA of ~2845 nt (EU273818 = NC_010417) (Yazdi et al. 2008; Heydarnejad et al. 2013; Soleimani et al. 2013; Varsani et al. 2014a; Zerbini et al. 2017).The genome encodes three predicted genes in the virion-sense, including the coat protein, and two predicted genes in the complementary sense. The replication associated protein is likely expressed from a spliced transcript, as is the case for geminiviruses of the genus Mastrevirus (Bozorgi et al. 2016). The genome of BCTIV also has an unusual nonanucleotide sequence, in the predicted hairpins structure which forms part of the origin of virion-strand DNA replication; “TAAGATTCC”' rather than the more usual “TAATATTAC” of other geminiviruses.

Beta vulgaris (Sugar beet)

Beet curly top virus

253

(BCTV)

Synonyms Beet mild curly top virus (BMCTV); Beet severe curly top virus (BSCTV)

B Taxonomic position Genus: Curtovirus

Family: Geminiviridae

Geographical distribution BCTV infection in plants of Beta vulgaris was reported from EPPO region (Cyprus, Egypt, Italy, Spain, Turkey), Asia (India, Iran, Turkey), Africa (Egypt), North America (British Columbia, Mexico, the USA), Central America and the Caribbean (Costa Rica, Puerto Rico), South America (Argentina, Bolivia, Brazil, Uruguay), and certain European countries (Bennett 1971; Magyarosy and Duffus 1977; Singh 1978; Duffus 1986; Polak 1988; Briddon et al. 1998; Wintermantel et al. 2003; Soto et al. 2005; Farzadfar et al. 2006; Strausbaugh et al. 2008, 2017; Lam et al. 2009; Anabestani et al. 2016; Strausbaugh et al. 2017). Some isolates of the virus have previously also been called Beet mild curly top virus, Beet severe curly top virus, Pepper yellow dwarf virus, and Spinach curly top virus (Baliji et al. 2004; Lam et al. 2009). Symptoms and host(s) The virus-infected sugar beet plants exhibit vein-clearing in young leaves that is followed by upward and inward leaf rolling. A conspicuous vein swelling and galling produce a roughened lower leaf surface, rasp-like to the touch. Droplets of phloem exudate often form on petioles and large veins. Virus infection induces increased number of rootlets and phloem necrosis observed as dark concentric rings in transverse sections of the roots. Older plants may turn yellow and die. The virus infects nearly 300 species of dicotyledonous plants of 44 families (Bennett 1971; Strausbaugh et al. 2008). On inoculation Spinacia oleracea, certain cucurbits, Phaseolus vulgaris, Linum spp., Capsicum spp., Solanum lycopersicum, Solanum tuberosum, and many other species exhibit vein-clearing; swelling and malformation in young leaves; rigid, dwarfed, yellowed, twisted, and malformed leaves; auxiliary buds stimulated; phloem necrosis; exudation of fluid from phloem; and death of the plant (Hosseini Abhari et al. 2005). Transmission The virus is transmitted by the leafhopper vector Circulifer tenellus in North America and C. tenellus and C. opacipennis in the Mediterranean basin in a circulative, non-propagative manner (Ball 1909; Soto and Gilbertson 2003; Rondon et al. 2016). The virus circulates but does not multiply in the vector and is not transmitted through the egg. Leafhoppers transmit occasionally after an acquisition time of 1 min, but the frequency of transmission is greatest after acquisition access periods of 2–3 days. The minimum latent period in the vector is rarely less than 4 h. The minimum probing period required for transmission is about 1 min. The frequency of transmission depends on the amount of virus in the vector which in turn depends on the virus content of the source plant. Highly infective vectors transmit the virus for life. The virus is not transmitted congenitally to the progeny of the vector (Bennett 1971; Thomas and Boll 1977). The virus is not transmitted by mechanical inoculation (unless special procedures are used), transmitted by grafting (by dodder from plants that are not hosts of the vector), and not transmitted by seed.

254

Beta vulgaris (Sugar beet)

Experimentally the virus has been introduced into sugar beet and Nicotiana benthamiana by Agrobacterium-mediated inoculation of cloned viral genomes and by needle inoculation with cloned viral DNA (Stanley et al. 1986; Briddon et al. 1989). Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein (Mumford 1974). The genome of BCTV consists of a single circular ssDNA molecule of ~2993 nt (M24597 = NC_001412) (Stanley et al. 1986; Stenger et al. 1990; Stenger 1994; Briddon et al. 1999; Teng et al. 2010; Varsani et al. 2014b; Zerbini et al. 2017). The genome encodes four complementary-sense genes and three virion-sense genes V1, V2, and V3. In the complementary-sense, these encode the replication-associated protein (Rep), the replication-enhancer protein (REn), and the C2 and C4 proteins (Stanley et al. 1986). The C2 protein has a role in pathogenicity in some hosts and suppresses transcriptional and posttranscriptional gene silencing (RNA interference-based host defense; Wang et al. 2005; Buchmann et al. 2009). The C4 protein plays a role in symptom development and cell-cycle control (Gutierrez 2002; Stanley 2008). BCTV C4 is responsible for tumorigenic growth in infected plants (Latham et al. 1997). In the virion-sense, the V1 and V3 gene products are required for the efficient movement of the virus in the infected plant (Hormuzdi and Bisaro 1993), whereas the coat protein interacts with the vector to mediate insect transmission (Briddon et al. 1990), is involved in virus movement in plants (Briddon et al. 1989), and may be involved in the regulation of ssDNA to dsDNA levels (Hormuzdi and Bisaro 1993; Briddon and Stenger 2011; Varsani et al. 2014b).

Beet leaf curl virus

(BLCV)

Taxonomic position BLCV is a tentative member of the family Rhabdoviridae. Geographical distribution BLCV infection in plants of Beta vulgaris was reported from the Eurasian region and the Mediterranean region, the former Czechoslovakia, Germany, Poland, Turkey, and the former USSR (Eisbein 1976; OEPP/EPPO 1982; Proeseler 1983). Symptoms and host(s) There is an initial glassiness or translucence of the leaf veins and petioles in virus-infected sugar beet leaves, which then become swollen and, since they do not grow as fast as the rest of the leaf, acquire a markedly crinkly appearance. The crown is stimulated to form new leaves which remain small and curve inward to form a compact bunch, resembling the head of cabbage lettuce. Growth is arrested; older leaves die. However, death of the plant before harvest rarely occurs. Transmission The virus is transmitted by the lace bug, Piesma quadratum, in the persistent (propagative) manner. The virus multiplies in its vector, and the bugs remain infective for life. There is no evidence of transmission to progeny insects. Minimum acquisition feeding period is 30 min; minimum inoculation feeding period is 40 min. There is a latent period of 7–35 days. Both the adult and nymphal stages can acquire and transmit the virus. The virus is not mechanically transmissible but has been transmitted by grafting to spinach (Spinacia oleracea) and Tetragonia tetragonioides (Schmutterer and Ehrhardt 1966).

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255

Virion properties and genome The virion morphology is bacilliform and the size is 80  225–350 nm (Proeseler 1983).

Beet mild yellowing virus Taxonomic position Genus: Polerovirus

(BMYV)

Family: Luteoviridae

Geographical distribution BMYV infection in plants of Beta vulgaris was reported from the Eurasian region, the British Isles, Russia, Spain, and the Western USA (Govier 1985; Stevens et al. 1995, 2004; Roman et al. 1996; Hauser et al. 2000; Mroczkowska et al. 2007). Symptoms and host(s) The virus-infected sugar beet plants exhibit symptoms of yellowing of older leaves, which become curled, thickened, and brittle with orange-yellow discoloration. Capsella bursa-pastoris, Stellaria media, Lamium amplexicaule, Senecio vulgaris, and Vernonia spp. are infected with this virus. Transmission The virus is transmitted by aphid vectors, Myzus persicae, M. (Sciamyzus) ascolonicus, M. (Nectarosiphon) certus, Macrosiphum euphorbiae, Acyrthosiphon (Aulacorthum) solani, Myzus (Phorodon) humuli, Brachycaudus helichrysi, and Aphis fabae, in a circulative, non-propagative manner (Govier 1985; Herrbach et al. 1992; Schliephake et al. 2000). The virus is retained when the vector molts and does not multiply in the vector and is not transmitted congenitally to the progeny of the vector. The virus is not transmissible by mechanical sap-inoculation, not by contact between plants, not by seed, and not by pollen. Virion properties and genome The virions are 25–30 nm in diameter, are hexagonal in outline, and have no envelope. The genome consists of a single molecule of positive-sense ssRNA. The complete nucleotide sequence of the genome of BMYV isolate 2ITB is 5722 nt (X83110). The 30 terminus has neither a poly(A) tract nor a tRNA-like structure. The 50 terminus has a genome-linked protein (VPg) (of about 17 kDa). The genome contains six open reading frames (ORFs) (Guilley et al. 1995; Stevens et al. 2005).

Beet mosaic virus Taxonomic position Genus: Potyvirus

(BtMV)

Family: Potyviridae

Geographical distribution BtMV was first reported in the late 1950s on Beta vulgaris from Germany (Russell 1971). The virus is distributed in all sugar beet-growing areas throughout the world, in particular in South Yugoslavia, Kazakhstan, Asia, China, Egypt, certain European countries, and the USA (Avgelis and Katis 1992; Rogov et al. 1992; Sutic et al. 1999; Glasa et al. 2000; Mali et al. 2000; Choueiri et al. 2001; Omar et al. 2005, 2006; Farzadfar et al. 2006; Xiang et al. 2007).

B

256

Beta vulgaris (Sugar beet)

Symptoms and host(s) The virus-infected sugar beet plants express symptoms of numerous small yellow spots and blotches on one or several central leaves. A light mosaic and mottle occurs on young leaves as the disease develops. Leaflets with initial symptoms are stunted, with curling and rolling of leaf margins and leaf tip necrosis. In severe cases, diseased leaves roll into a tubular shape (Wintermantel 2005). The natural host range of this virus includes Beta maritima, Melilotus indicus, Trifolium incarnatum, Spinacia oleracea, Chenopodium album, Amaranthus retroflexus, and Sonchus arvensis. Transmission The virus is transmitted by more than 28 aphid spp. including Myzus persicae, Aphis fabae, Rhopalosiphum padi, Acyrthosiphon (Metopolophium) dirhodum, and Macrosiphum (Sitobion) avenae. Principal natural vectors include Myzus persicae and Aphis fabae. The virus-vector relationship is of the non-persistent type (Dusi and Peters 1999). The virus is transmissible by mechanical sapinoculation to cultivated sugar beet and also to a large number of herbaceous plants belonging to the families Chenopodiaceae, Solanaceae, and Leguminosae. The virus is transmissible by grafting, not transmissible by contact between plants, and not transmissible by pollen (Dusi and Peters 1999). Virion properties and genome The virions are non-enveloped, flexuous filamentous particles of 680–900 nm long and 11–13 nm wide. Virus particles are coated with approximately 2000 copies of a single capsid protein; the molecular weight of each capsid protein subunit is about 34,700 Da (Rogov et al. 1991). The genome consists of single molecule of linear positive-sense ssRNA. The complete genome of BtMV-Wa isolate from the USA is 9591 nt (AY206394 = NC_005304) long with a poly(A) tail at the 30 end (Nemchinov et al. 2004; Revers and Garcia 2015; Wylie et al. 2017).

Beet necrotic yellow vein virus Taxonomic position Genus: Benyvirus

(BNYVV)

Family: Benyviridae

Geographical distribution BNYVV in Beta vulgaris occurs in almost all countries, viz., Austria, Belgium, Bulgaria, Croatia, the Czech Republic, Brazil, France, Germany, Greece, Hungary, Italy, the Netherlands, Poland, Romania, Slovakia, Slovenia, Spain, Sweden, Switzerland, the UK, Russia, Turkey, Yugoslavia, China, Japan, Kazakhstan, Kyrgyzstan, Mongolia, multiple states of the USA and many European countries as well as the USSR (Duffus et al. 1984; Lesemann et al. 1989; Whitney and Duffus 1991; Vardar and Erkan 1992; Asher 1993; Konecny 1994; Choueiri et al. 2001; Kutluk and Yanar 2001; Nielsen et al. 2001; Tamada 2001; Mouhanna et al. 2002; Meunier et al. 2003; Kutluk et al. 2004; Farzadfar et al. 2007; Yardimci and Culal Kilic 2011; Zizyte et al. 2013; Rezende et al. 2015; Ebrahim-Ghomi et al. 2016; Kutluk et al. 2016). From Poland and Sweden, type A and B of BNYVV in sugar beets have been reported (Lennefors et al. 2000; Borodynko 2006a; Borodynko et al. 2009; Roberts et al. 2016). Three distinct genotypes of BNYVV, A, B, and P type, have been identified in Europe which mostly occur in separate geographic areas (Koenig and Lennefors 2000; Ward et al. 2007). The A- and B-type infections are most common. The B type is prevalent in Central Europe, especially in Germany and France. The A type is almost exclusively distributed in Southern Europe; however, it is also distributed in countries along the Atlantic Coast (Belgium, the Netherlands) as well as in England, Sweden, and several Eastern European countries (Koenig and Lennefors 2000; Ratti et al. 2005; Schirmer et al.

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2005). The P type has so far been found around the French town of Pithiviers (Koenig et al. 1995) and in limited areas in the UK (Ward et al. 2007).

Symptoms and host(s) BNYVV causes rhizomania disease of sugar beet (Canova 1959; Tamada and Baba 1973; Putz et al. 1990; Richards and Tamada 1992). The most characteristic symptom is root stunting and proliferation of the lateral roots which are dark brown bearded and pale yellow to dark brown vascular discoloration in transverse section. In early infections, plants are often stunted and can wilt and die; in later infections, the virus may reach the leaves, causing yellow and necrotic vein symptoms. Other characteristic symptoms are elongation of leaf petioles and narrowing of leaf blades. The narrower leaf blades are erect in the center of the plant and thus can easily be observed. The host range is very narrow and infects all subspecies of Beta vulgaris, including sugar beet (Beta vulgaris sub spp. maritima), fodder beet (Beta vulgaris sub spp. vulgaris), red beet (Beta vulgaris sub spp. cicla), mangolds (Beta vulgaris sub spp. vulgaris), sea kale (Beta vulgaris sub spp. vulgaris), Swiss chard (Beta vulgaris sub spp. cicla), and also spinach (Spinacea oleracea) and species of Chenopodiceae (Tamada 2002).

Transmission The virus is transmitted by the fungal vector, Polymyxa betae, belonging to the Plasmodiophoraceae (Tamada et al. 1975; Fujisawa and Sugimoto 1976; Peters and Godfrey-Veltman 1989; Abe and Tamada 1986; Hugo et al. 1996; Meunier et al. 2003; Kutluk et al. 2010). The long persistence of P. betae resting spores carrying the virus in the soil allows infection of sugar beet more than 10 years after the last crop. P. betae is able to survive in the soil by means of thick-walled resting spores, named sporosori, released during root decay. These sporosori are able to survive in the soil for years. Under favorable conditions, the sporosori will evolve into zoospores which cause fresh infections in humid soils. The virus is not transmitted by Polymyxa graminis, Myzus persicae, and Acyrthosiphon (Aulacorthum) solani. The virus is transmissible by mechanical sap-inoculation, not transmissible by contact between plants, not transmitted by seed, and not transmitted by pollen (Stevens et al. 2006).

Virion properties and genome The virions are non-enveloped, helically constructed rod-shaped particles, with an axial canal and of up to five different lengths are about 390, 265, 100, 85 and 65-80 nm and their diameter is about 20 nm. The right-handed helix with a pitch of 2.6 nm has an axial repeat of four turns, involving 49 CP subunits, each occupying four nucleotides (Steven et al. 1981). Coat proteins constitute about 95% of the particle weight. BNYVV is a multicomponent positive-sense ssRNA virus. BNYVV contains 4–5 single-stranded capped (m7G) and polyadenylated RNAs (Putz 1977; Putz et al. 1983). The RNA-1 (D84410 = NC_003514) is 6746 nt excluding the poly(A) tail and has one long open reading frame (ORF) encoding a 237.34 kDa replicase-associated protein. The RNA-2 (D84411 = NC_003515), RNA-3 (D84412 = NC_003516), RNA-4 (D84413 = NC_003517) and RNA-5 (D63936 = NC_003513) are 4609, 1774, 1465 and 1320 nt, respectively (Bouzoubaa et al. 1985, 1986, 1987, 1991; Kiguchi et al. 1996; Saito et al. 1996; et al. 1997a, 1999, 2009; Tamada et al. 1996; Harju et al. 2002; Delbianco et al. 2013; Kutluk et al. 2007; Ward et al. 2007; Gilmer et al. 2011, 2017).

B

258

Beta vulgaris (Sugar beet)

Beet pseudoyellows virus Taxonomic position Genus: Crinivirus

(BPYV)

Family: Closteroviridae

Geographical distribution BPYV infection in plants of Beta vulgaris was first reported from Salinas, California, USA, by Duffus (1965). Subsequently it was also recorded in Australia, France, Belgium, Italy, Turkey, Japan, and the Netherlands (Polak and Lot 1988; Rubio et al. 1999; Tzanetakis et al. 2013; Abrahamian and AbouJawdah 2014). Symptoms and host(s) The virus-infected sugar beet plants exhibit interveinal chlorosis, yellowing, thickening, and brittleness of the leaves. The virus has a broad host range including other crops (lettuce, spinach, endive), ornamentals (marigold, zinnia), and weeds (dandelion, cheeseweed) (Duffus 1965; Wisler et al. 1998). Transmission The virus is transmitted by a whitefly vector, Trialeurodes vaporariorum, in a semi-persistent manner (Duffus 1965; Wisler et al. 1998; Jones 2003). Through the insect vector the virus infects more than nine plant families. The virus does not multiply in the vector and is not transmitted congenitally to the progeny of the vector. The virus is not transmissible by mechanical inoculation, not transmissible by contact between plants, and not transmissible by seed. Virion properties and genome The virions are non-enveloped, bipartite filamentous particles about 650–850 nm and 700–900 nm in length, and 10–13 nm in diameter (Liu and Duffus 1990). The genome consists of two molecules of positive-sense ssRNA. The RNA-1 genome segment was 8006 nt containing two ORFs (ORF-1a and ORF-1b and ORF-2) (AY330918 = NC_005209). The RNA-2 genome segment is 7903 nt containing seven ORFs (AY330919 = NC_005210) (Kreuze 2011; Tzanetakis et al. 2013).

Beet ringspot virus

(BRSV)

Taxonomic position Genus: Nepovirus

Family: Secoviridae

Geographical distribution BRSV infection in plants of Beta vulgaris was reported from Scotland (Harrison 1957). Symptoms and host(s) The virus-infected sugar beet plants show necrotic ringspots, systemic chlorotic ringspots, mottling, and vein yellowing symptoms. Transmission The virus is transmitted efficiently by the soil-borne nematode vector, Longidorus elongatus, in a non-persistent manner. In glasshouse experiments sugar beet, beetroot, potato, turnip, swede, French bean, Fragaria vesca, and oat and wheat plants often became systemically infected when grown in soil

Beta vulgaris (Sugar beet)

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from the site of a disease outbreak, but the virus was restricted to the roots of many infected plants. The virus is mechanically sap-transmissible (Harrison 1957). Virion properties and genome The virions are isometric, non-enveloped of two types, but similar in size of 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is bipartite, linear, positive-sense, single-stranded RNA. The RNA-1 genome segment is 7362 nt that contains a single ORF encoding a 254 kDa polyprotein (D00322 = NC_003693) (Greif et al. 1988). The RNA-2 is 4662 nt (X04062 = NC_003694) long encoding a polyprotein (Meyer et al. 1986). Movement protein and coat protein are located in the RNA-2 genome segment (Sanfacon et al. 2009; Dunez and Le Gall 2011; Sanfacon 2015; Thompson et al. 2017).

Beet soil-borne mosaic virus Taxonomic position Genus: Benyvirus

(BSBMV)

Family: Benyviridae

Geographical distribution BSBMV infection in plants of Beta vulgaris was reported from Iran, China, and the USA (Rush and Heidel 1995; Heidel et al. 1997; Rush et al. 1997; Wisler et al. 1997b). Symptoms and host(s) Foliar symptoms produced by this virus include slight leaf distortion, subtle overall mottling, and light green or yellow blotches and bands that follow primary leaf veins. As diseased leaves age, the bands can progress to broad chlorotic areas that usually remain associated with the veins. Occasionally, systemically infected leaves exhibit a mottled or mosaic pattern or symptoms. Some BSBMV isolates cause bright yellow veinal chlorosis. Roots are often asymptomatic, but some beets systemically infected with BSBMV exhibit root symptoms more typically associated with rhizomania, including stunting, proliferation of lateral roots, and constriction of the main taproot. Transmission The virus is transmitted by the soil-borne fungus Polymyxa betae (Wisler et al. 2003). The virus is also transmissible by mechanical sap-inoculation. Virion properties and genome The virions are rigid, rod-shaped particles of 50–400  19 nm. The molecular mass of the capsid protein is 22.5 kDa (Heidel and Rush 1997; Heidel et al. 1997; Lee et al. 2001). The virus is a multicomponent positive-sense ssRNA virus. The genome consists of four RNA segments: RNA-1 consists of 6679 nt (JF513082; AF280539), RNA-2 of 4615 nt (JF513083; AF061869), RNA-3 of 1720 nt (EU410955; AF280540), and RNA-4 of 1730 nt (FJ424610; AF280541). The RNA-1 genome segment contains a single large open reading frame (ORF) encoding replicase-associated protein similar to BNYVV. The RNA-2 contains six ORFs with an organization resembling BNYVV RNA-2 (Lee et al. 2001; Gilmer et al. 2011, 2017).

B

260

Beta vulgaris (Sugar beet)

Beet soil-borne virus Taxonomic position Genus: Pomovirus

(BSBV)

Family: Virgaviridae

Geographical distribution BSBV infection in plants of Beta vulgaris was first described in England (Henry et al. 1986) and has since been found in all sugar beet-growing areas worldwide including Europe, China, Greece, Iran, the Netherlands, Belgium, Sweden, Germany, France, Turkey, and the USA (Verhoyen et al. 1987; Lindsten 1989; Heidel et al. 1993; Lindsten and Rush 1994; Farzadfar et al. 2002b, 2007; Meunier et al. 2003; Kutluk et al. 2004; Borodynko et al. 2006; Wang et al. 2008a; Pavli et al. 2010; Kutluk et al. 2010, 2016). Symptoms and host(s) Sugar beet is frequently a symptomless host but cultivar Monoire exhibited chlorotic rings (Prillwitz and Schlosser 1992). Transmission The virus is transmitted by a fungal vector, Polymyxa betae (Meunier et al. 2003). The virus is transmitted by mechanical sap-inoculation, and inoculated Chenopodium quinoa and C. amaranticolor leaves produced local lesions (Wang et al. 2007). The possibility of transmission by contact between plants (but not easily) was reported by Ivanovic et al. (1983). Virion properties and genome The virions are rod-shaped particles that have predominant lengths of about 65, 150, and 300 nm and diameters of 19 nm (Koenig et al. 1997b). The virus contains three segments of linear positive-sense ssRNA. The complete genome sequence of BSBV Ahlum isolate has been determined. The RNA segments are capped at 50 and folded into a tRNA-like structure at the 30 end. The RNA-1 genome segment is 5834 nt (Z97873 = NC_003520) containing a readthrough protein of 204 kDa, which is interrupted resulting in the formation of a protein of 145 kDa. The RNA-2 genome segment contains 3454 nt (U64512 = NC_003518) (Koenig et al. 1997b). The RNA-3 genome segment is 3005 nt (Z66493 = NC_003519) and is unique among the plant virus RNAs as it encodes only a triple gene block (TGB) (Koenig et al. 1996; Savenkov 2011; Adams et al. 2017).

Beet virus Q Taxonomic position Genus: Pomovirus

(BVQ)

Family: Virgaviridae

Geographical distribution BVQ infection in plants of Beta vulgaris was reported from Greece, Germany, Belgium, Bulgaria, France, Hungary, Iran, Italy, Poland, Spain, Sweden, and the Netherlands (Stas et al. 2001; Meunier et al. 2003; Farzadfar et al. 2005, 2007; Borodynko 2006b; Rubies Autonell et al. 2006; Pavli et al. 2010).

Beta vulgaris (Sugar beet)

261

Symptoms and host(s) The virus-infected sugar beet plants produce yellowing of the leaves and also rhizomania-like symptoms on the roots. Transmission The virus is transmitted by a plasmodiophoromycete fungal vector, Polymyxa betae (Meunier et al. 2003; Farzadfar et al. 2007). The mechanically inoculated sugar beet plants produces irregularly shaped local lesions after 5 days that tend to spread along the veins. Mechanically inoculated Chenopodium quinoa and C. amaranticolor leaves produced local lesions. The virus could not be transmitted to Nicotiana benthamiana or Nicotiana clevelandii. Virion properties and genome The virions are rod-shaped particles that have predominant lengths of about 65–80, 150–160, and 290–310 nm and diameters of 18–20 nm. The genome consists of three segments of linear positivesense ssRNA. RNA-1 consists of 6003 nt (AJ223596 = NC_003510), RNA-2 of 2913 nt (AJ223597 = NC_003511), and RNA-3 of 2529 nt (AJ223598 = NC_003512) (Koenig et al. 1998; Savenkov 2011; Adams et al. 2017).

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

Geographical distribution BWYV infection in plants of Beta vulgaris is probably distributed worldwide (Marco 1984; Smith and Hinckes 1985; Stevens et al. 1995;Choueiri et al. 2001; Farzadfar et al. 2006; Beuve et al. 2008; Xiang et al. 2008; Asaad et al. 2009; Zhou et al. 2011). Symptoms and host(s) The virus-infected sugar beet plants exhibit mild chlorotic spotting, yellowing, thickening, and brittleness of older leaves. Certain beet cultivars show an excessive proliferation of nonfunctional, necrotizing secondary rootlets (rhizomania) (Sanger et al. 1994; Wintermantel 2005). More than 150 species in 23 dicotyledonous families are susceptible to the virus including such economically important species as Spinacia oleracea, Helianthus annuus, Lactuca sativa, Brassica napobrassica, B. napus, B. nigra, B. oleracea var. botrytis and capitata, B. rapa, Raphanus sativus, Crambe abyssinica, Citrullus lanatus, Cucumis sativus, Cucurbita pepo, Cicer arietinum, Glycine max, Pisum sativum, Trifolium subterraneum, Vicia faba, Phlox drummondii, Capsicum annuum, and Solanum lycopersicum (Timmerman et al. 1985; Coutts et al. 2006; Asaad et al. 2009). Transmission The virus is transmitted by aphid vectors, Myzus persicae, Aphis craccivora, A. gossypii, Acyrthosiphon (Aulacorthum) solani, Brachycaudus helichrysi, Brevicoryne brassicae, Macrosiphum euphorbiae, Myzus (Sciamyzus) ascalonicus, M. ornatus, and Myzus (Phorodon) humuli, in a circulative, non-propagative manner. The virus is retained when the vector molts, does not multiply in the vector, and is not transmitted congenitally to the progeny of the vector, not transmissible by mechanical inoculation, not transmissible by grafting, not transmissible by contact between plants, not transmissible by seed, and not transmissible by pollen.

B

262

Beta vulgaris (Sugar beet)

Virion properties and genome The virions are 25–30 nm in diameter, are hexagonal in outline, and have no envelope. The genome consists of a single molecule of positive-sense ssRNA. The genome of BWYV lettuce isolate is 5641 nt (AF473561). The 30 terminus has neither a poly(A) tract nor a tRNA-like structure. The 50 terminus has a genome-linked protein (VPg) (of about 17 kDa). The genome contains six open reading frames (ORFs) (Veidt et al. 1988; Pfeffer et al. 2002).

Beet yellow stunt virus Taxonomic position Genus: Closterovirus

(BYSV)

Family: Closteroviridae

Geographical distribution BYSV infection in plants of Beta vulgaris was reported from the USA (California) (Hoefert et al. 1970; Duffus 1972). Symptoms and host(s) In the virus-infected sugar beet plants, initially there is a severe twisting and cupping of leaves that become mottled and yellow. Diseased plants are severely stunted. Lactuca sativa and Sonchus oleraceus are susceptible. Transmission The virus is transmitted by aphid vectors, viz., Hyperomyzus lactucae, Myzus persicae, Macrosiphum euphorbiae, in a semi-persistent manner. The principal natural vector is Hyperomyzus lactucae. The virus is lost by the vector when it molts and not transmitted congenitally to the progeny of the vector. The virus is not transmissible by mechanical inoculation, not transmissible by contact between plants, and not transmissible by seed. Virion properties and genome The virions are long, flexuous filaments rods approximately 1400 nm long and 12.5 nm in width. The BYSV coat protein of 23.7 kDa is expressed from an approximately 1.9-kb sgRNA (Karasev et al. 1998). The genome consists of a single molecule of large positive-sense ssRNA of about 18,000 nt (Reed and Falk 1989; Karasev et al. 1998). The 30 -terminal half of the BYSV genome of 10,545 nt (U51931) has been determined that contains ten open reading frames (ORFs) and 241 nt of the 30 untranslated region (Karasev et al. 1996). The five-gene module conserved among closteroviruses was identified in BYSV, which includes a gene array coding for a 6-kDa small hydrophobic protein, a 66-kDa homolog of the cellular HSP70 heat shock proteins, a 61-kDa protein, and a 25-kDa diverged copy of the CP followed by the CP gene itself (Agranovsky and Lesemann 2011).

Beet yellows virus Taxonomic position Genus: Closterovirus

(BYV)

Family: Closteroviridae

Beta vulgaris (Sugar beet)

263

Geographical distribution BYV is widespread wherever Beta vulgaris is grown, in Europe, Asia, North and South America, Africa, Australia, Spain, Croatia, Lebanon and China (Roland 1936; Bennett 1960; Shepherd and Hills 1970; Roman et al. 1996; Choueiri et al. 2001; Wintermantel 2005; Farzadfar et al. 2006; Voncina et al. 2015).

Symptoms and host(s) The virus-infected sugar beet show vein yellowing or vein-clearing in younger leaves. Leaves turn chlorotic, notably thickened, leathery, and brittle. Reddish spots appear between leaf veins, grow larger, and often merge. The leaves may have a distinct bronze cast. Infected plants often turn reddish or purplish, or the normal reddish or purplish color intensifies. Finally, about the time seeds are maturing, the plant is in general decline. Affected seed plants have poor root growth and are easily uprooted by a moderate wind. Their seed is smaller and has less vitality than seed from uninfected plants.

Transmission The virus is transmitted by 35 aphid species of which Myzus persicae and Aphis fabae are the most important vectors and transmit the virus in a semi-persistent manner (Heathcote and Cockbain 1964; Limburg et al. 1997). The virus is lost by the vector when it molts, does not multiply in the vector, and not transmitted congenitally to the progeny of the vector. On the other hand, Rhopalosiphum padi and Macrosiphum (Sitobion) avenae failed to transmit this disease. The virus is transmissible by mechanical sap-inoculation (with difficulty). Nearly 130 species are susceptible to this virus of which 23 are natural hosts (Schmelzer and Hartleb 1977). The virus is also transmissible by grafting but not transmissible by contact between plants, not transmissible by seed, and not transmissible by pollen.

Virion properties and genome The virions are filamentous, non-enveloped, very flexuous, 1250 nm long, and 10 nm in diameter. The virion is composed of a long body and short tail whose principal components are the major and minor capsid proteins, respectively. BYV CP, CPm, p64, and Hsp70h are each required for virion assembly (Peremyslov et al. 2004). The virus has a single molecule of linear, positive-sense ssRNA genome of 15,468 nt containing 8 open reading frames (ORFs) (AF190581, X73476). The 50 genome region contains two overlapping open reading frames (ORF)-1a and ORF-1b encoding 295 K and 48 K proteins, respectively. The RNA sequence surrounding the stop codon in ORF-1a shows structural elements typical of ribosomal frameshifting signals in a number of animal and plant viruses. It is predicted that the ORF-1b product is expressed via a +1 ribosomal frameshifting as the 348 K ORF-1a/ORF-1b fusion protein. This putative protein contains the array of methyl transferase, RNA helicase, and RNA-dependent RNA polymerase domains that are conserved in the Sindbis-like super group of positive-strand RNA viruses (Agranovsky et al. 1994). The 30 -terminal portion of the BYV genome contains seven ORFs. In the replication-associated 1a poly protein of BYV, the region between the methyl transferase and helicase domains (1a central region (CR), 1a CR) is marginally conserved. The CR-2 may be involved in the formation of multivesicular complexes in BYV-infected cells (Agranovsky and Lesemann 2011; Gushchin et al. 2013).

B

264

Beta vulgaris (Sugar beet)

Chickpea chlorotic dwarf virus Taxonomic position Genus: Mastrevirus

(CpCDV)

Family: Geminiviridae

CpCDV infection in plants of Beta vulgaris was reported from Iran (Farzadfar et al. 2002a, 2006, 2008). The virus-infected sugar beet plants exhibit chlorosis and stunting. The virus is transmitted by the leafhopper vector Orosius orientalis in a persistent, circulative, and non-propagative manner, and infects a number of economically important crops. The virus is not transmitted by mechanical inoculation. For more details of CpCDV, refer to Cicer arietinum.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV is distributed wherever Beta vulgaris is grown (Alhubaishi et al. 1987; Farzadfar et al. 2006). The virus-infected sugar beet plants exhibit large, irregular-shaped patches of bright yellow tissue which contrast with the dark green of the leaf. Symptoms are of a mosaic type, but the pattern is coarser and more contrasting than that found in beet mosaic. The leaves commonly are dwarfed and distorted and have blister-like areas of green tissue. Necrosis often occurs in large areas of the diseased older leaves. The virus is transmitted by more than 60 aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Lettuce chlorosis virus Taxonomic position Genus: Crinivirus

(LCV)

Family: Closteroviridae

LCV occurs on Beta vulgaris in the USA (Duffus et al. 1996; Wisler et al. 1997a). The virus-infected sugar beet plants exhibit interveinal yellowing and thickened leaves. Interveinal reddening occurs in the leaves of inoculated plants. The virus is transmitted by the sweet potato whitefly Bemisia tabaci “B” biotype in a semi-persistent manner. The virus is not mechanically sap-transmissible. For more details of LCV, refer to Lactuca sativa.

Lettuce infectious yellows virus Taxonomic position Genus: Crinivirus

(LIYV)

Family: Closteroviridae

LIYV occurs on Beta vulgaris in California (USA) (Duffus et al. 1986). The virus-infected sugar beet plants show very mild mottling that develops into an interveinal yellowing of diseased leaves. A necrosis eventually develops in the chlorotic areas of the diseased leaves. The virus is transmitted by the sweet potato whitefly, Bemisia tabaci, in a semi-persistent manner. The virus is not mechanically sap-transmissible. For more details of LIYV, refer to Lactuca sativa.

Beta vulgaris (Sugar beet)

265

Soybean dwarf virus

(SbDV)

Synonyms Subterranean clover red leaf virus

B Taxonomic position Genus: Luteovirus

Family: Luteoviridae

SbDV infection in plants of Beta vulgaris was reported from northwestern Tasmania, Europe, and North America (Johnstone and Munro 1980). The virus-infected sugar beet plants showed the initial symptom of interveinal chlorosis on the older leaves which developed to a bright leaf yellowing which was occasionally accompanied by a veinal necrosis. Yellowed leaves sometimes developed orange tints. Symptoms rarely extended to the younger leaves, but these occasionally had a mottled appearance on severely affected plants. Leaves with symptoms generally were thickened and brittle. The virus is transmitted by the aphid vector Aulacorthum solani in a persistent manner. The virus is not mechanically sap-transmissible. For more details of SbDV, refer to Glycine max.

Tobacco mosaic virus

(TMV)

Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

TMV infection in Beta vulgaris plants was reported from China (Krstic et al. 1997; Wang et al. 2008b). The virus is mechanically sap-transmissible and also through contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco rattle virus

(TRV)

Taxonomic position Genus: Tobravirus

Family: Virgaviridae

TRV infection in plants of Beta vulgaris was reported from Bulgaria (Dikova 2006). The virus-infected sugar beet plants exhibit yellowing and chlorotic mottling symptoms. The virus is transmitted by nematode vectors and also by mechanical sap-inoculation to a large number of host plants. The virus is seedtransmitted up to 15–20% in sugar beet (Dikova 2005). For more details of TRV, refer to Nicotiana tabacum.

Tomato black ring virus Taxonomic position Genus: Nepovirus

(TBRV)

Family: Secoviridae

TBRV infection in plants of Beta vulgaris was reported from the UK (CABI 2015). The virus-infected sugar beet plants exhibit ringspot symptoms. The virus is transmitted by the nematode vector

266

Beta vulgaris (Sugar beet)

Longidorus attenuatus in a non-persistent manner, and also by mechanical sap-inoculation (Brown and Trudgill 1998). For more details of TBRV, refer to Solanum lycopersicum.

Tomato bushy stunt virus Taxonomic position Genus: Tombusvirus

(TBSV)

Family: Tombusviridae

TBSV infection in plants of Beta vulgaris was reported from Czechoslovakia (Novak and Lanzova 1980). The virus-infected sugar beet plants show large chlorotic ringspots and line pattern. The virus is transmitted through the soil although the biological vector is not confirmed. The virus is mechanically sap-transmissible and infects a large number of hosts. For more details of TBSV, refer to Solanum lycopersicum.

Turnip curly top virus Taxonomic position Genus: Turncurtovirus

(TCTV)

Family: Geminiviridae

TCTV infection in plants of Beta vulgaris was reported from Iran (Kamali et al. 2016). The virus is transmitted by the leafhopper vector Circulifer haematoceps in a circulative and non-propagative manner. For more details of TCTV, refer to Brassica rapa.

Turnip leaf roll virus Taxonomic position Genus: Turncurtovirus

(TuLRV)

Family: Geminiviridae

TuLRV infection in plants of Beta vulgaris was reported from Iran (Kamali et al. 2016). The virusinfected sugar beet plants exhibit symptoms of inward curling of leaves. The virus is transmitted by the leafhopper vector Circulifer haematoceps in a circulative, non-propagative manner. For more details of TuLRV, refer to Brassica rapa.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Beta vulgaris was reported from Iran (Farzadfar et al. 2006). The virusinfected sugar beet plants exhibit mosaic, mottling, and vein-clearing symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

Beta vulgaris (Sugar beet)

267

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Stas A, Meunier A, Schmit J-F, Bragard C (2001) First report of Beet virus Q in Belgium. Plant Dis 85:1288 Stenger DC (1994) Complete nucleotide sequence of the hypervirulent CFH strain of Beet curly top virus. Mol PlantMicrobe Interact 7:154–157 Stenger DC1, Carbonaro D, Duffus JE (1990) Genomic characterization of phenotypic variants of Beet curly top virus. J Gen Virol 71:2211–2215 Steven AC, Trus BL, Putz C, Wurtz M (1981) The molecular organization of beet necrotic yellow vein virus. Virology 113(2): 428–438 Stevens M, Smith HG, Hallsworth PB (1995) Detection of the luteoviruses, Beet mild yellowing virus and Beet western yellows virus, in aphids caught in sugar-beet and oilseed rape crops, 1990-1993. Ann Appl Biol 127:309–320 Stevens M, Hallsworth PB, Smith HG (2004) The effects of Beet mild yellowing virus and Beet chlorosis virus on the yield of UK field-grown sugar beet in 1997, 1999 and 2000. Ann Appl Biol 144:113–120 Stevens M, Freeman B, Liu H-Y, Herrbach E, Lemaire O (2005) Beet poleroviruses: close friends or distant relatives? Mol Plant Pathol 6:1–9 Stevens M, Lui H-Y, Lemaire O (2006) The viruses. In: Draycott AP (ed) The sugar beet book. Blackwells, Oxford, UK, pp 256–285 Strausbaugh CA, Wintermantel WM, Gillen AM, Eujayl IA (2008) Curly top survey in the western United States. Phytopathology 98:1212–1217 Strausbaugh CA, Eujayl IA, Wintermantel WM (2017) Beet curly top virus strains associated with sugar beet in Idaho, Oregon, and a Western U.S collection. Plant Dis 101:1373–1382 Sutic DD, Ford RF, Tosic MT (1999) Beet mosaic potyvirus (BtMV). In: Handbook of plant virus diseases. CRC Press, Washington, DC, pp 279–280 Szego A (2009) Distribution, persistence and molecular characterization of cryptic and Endornaviruses. Thesis of doctoral dissertation, Corvinus University of Budapest. Budapest Szego A, Enunlu N, Deshmukh SD, Veliceasa D, Hunyadi-Gulyas E, Kühne T, Ilyes P, Potyondi L, Medzihradszky K, Lukacs N (2010) The genome of Beet cryptic virus 1 shows high homology to certain Cryptoviruses present in phylogenetically distant hosts. Virus Genes 40:267–276 Taheri H, Izadpanah K, Behjatnia SAA (2012) Circulifer haematoceps, the vector of the Beet curly top Iran virus. Iran J Plant Pathol 48:151 Tamada T (2001) Studies on Beet necrotic yellow vein virus. J Gen Plant Pathol 67:248–250 Tamada T (2002) Beet necrotic yellow vein virus. Association of Applied Biologists Descriptions of Plant Viruses, No. 144 Tamada T, Baba T (1973) Beet necrotic yellow vein virus from rhizomania-affected sugar beet in Japan. Ann Phytopathol Soc Jpn 39:325–332 Tamada T, Abe H, Baba T (1975) Beet necrotic yellow vein virus and its relation to the fungus Polymyxa betae. In: Proceedings of the first intersectional congress of international association of microbiological societies, vol 3. pp 313–320 Tamada T, Schmitt C, Guilley H, Richards K, Jonard G (1996) High resolution analysis of the read through domain of Beet necrotic yellow vein virus readthrough KTER motif is important for efficient transmission of the virus by Polymyxa betae. J Gen Virol 77:1359–1367 Teng K, Chen H, Lai J, Zhang Z, Fang Y, Xia R, Zhou X, Guo H, Xie Q (2010) Involvement of C4 protein of Beet severe curly top virus (Family Geminiviridae) in virus movement. PLoS One 5(6):e11280 Thomas PE, Boll RK (1977) Effect of host preference on transmission of curly top virus to tomato by the beet leafhopper. Phytopathology 67:903–905 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Secoviridae. J Gen Virol 98:529–531 Timmerman EL, D'Arcy CJ, Splittstoesser WE (1985) Beet western yellows virus in Illinois vegetable crops and weeds. Plant Dis 69:933–936 Tzanetakis IE, Martin RR, Wintermantel WM (2013) Epidemiology of criniviruses: an emerging problem in world agriculture. Front Microbiol 4:119 Vainio EJ, Chiba S, Ghabrial SA, Maiss E, Roossinck M, Sabanadzovic S, Suzuki N, Xie J, Nibert M, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Partitiviridae. J Gen Virol 99:17–18 Vardar B, Erkan S (1992) The first studies on the detection of Beet necrotic yellow vein virus in sugar beet Turkey. J Turk Phytopathol 21:71–79 Varsani A, Navas-Castillo J, Moriones E, Hernandez-Zepeda C, Idris A, Brown JK, Zerbini FM, Martin DP (2014a) Establishment of three new genera in the family Geminiviridae: Becurtovirus, Eragrovirus and Turncurtovirus. Arch Virol 159:2193–2203 Varsani A, Martin DP, Navas-Castillo J, Moriones E, Hernández-Zepeda C, Idris A, Murilo Zerbini F, Brown JK (2014b) Revisiting the classification of curtoviruses based on genome-wide pairwise identity. Arch Virol 159(7):1873–1882

B

274

Beta vulgaris (Sugar beet)

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Beta vulgaris var. cicla (Spinach beet/ Swiss chard)

275

Beta vulgaris var. cicla (Spinach beet/ Swiss chard) Family: Amaranthaceae

Leafy vegetable

B Beet cryptic virus 3 Taxonomic position Genus: Deltapartitivirus

(BCV-3)

Family: Partitiviridae

Geographical distribution BCV-3 was first reported in Beta vulgaris var. cicla from Italy in plants grown from Japanese seed (Xie et al. 1993). The virus spreads in Japan, Italy, and the UK. Symptoms and host(s) No symptoms are produced by BCV-3 on spinach beet. Transmission No vector is reported for this virus. The virus is not mechanically sap-transmissible and also not transmissible by grafting. The virus spreads through true seed collected from infected plants. Virion properties and genome The virions are isometric, non-enveloped, and 30 nm in diameter. BCV-3 contains two dsRNA genome components of 1740 bp and 1600 bp. The nucleotide sequence of BCV-3 RNA2 has been determined, which is 1607 bp (S63913) (Xie et al. 1993; Vainio et al. 2018).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Beta vulgaris var. cicla was reported from Croatia (Krajacic and Juretic 1993). The virus-infected spinach beet plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

References Krajacic M, Juretic N (1993) Natural infection of Swiss chard by Cucumber mosaic virus in Croatia. Petria 3:93–98 Xie WS, Antoniw JF, White RF (1993) Nucleotide sequence of Beet cryptic virus 3 dsRNA2 which encodes a putative RNA-dependent RNA polymerase. J Gen Virol 74:1467–1470 Vainio EJ, Chiba S, Ghabrial SA, Maiss E, Roossinck M, Sabanadzovic S, Suzuki N, Xie J, Nibert M, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Partitiviridae. Journal of General Virology 99:17–18

276

Betula spp. (Birch)

Betula spp. (Birch) Family: Betulaceae

Trees/Shrubs

Apple mosaic virus

(ApMV)

Taxonomic position Genus: Ilarvirus

Family: Bromoviridae

ApMV infection in plants of Betula spp. was reported from Finland, Eastern Canada, and Northeastern USA (Gotlieb and Berbee 1973; Gotlieb 1975; Cooper and Massalski 1984; Gruntzig et al. 1996; Polak and Prochazkova 1996; von Bargen et al. 2009). The virus-infected birch plants exhibit symptoms of chlorotic lines forming “oak leaf” pattern, irregular rings or linear flecks, and mild mosaic. Symptoms usually restricted to a few fully expanded leaves on some branches. No insect vector is known for this virus. The virus is mechanically sap-transmissible. For more details of ApMV, refer to Malus domestica.

Birch leaf roll-associated virus Taxonomic position Genus: Badnavirus

(BLRaV)

Family: Caulimoviridae

Geographical distribution BLRaV infection in plants of Betula pendula was reported from Germany (Rumbou et al. 2018). Transmission The virus is transmitted by mealybug vectors in a semipersistent manner. The virus is transmitted by mechanical inoculation (with difficulty). The virus is transmitted by grafting and not by contact between plants or by pollen. Virion properties and genome The virions are bacilliform in shape with a diameter of 30 nm and modal particle length of 130 nm. The genome is a circular dsDNA of 7863 bp (MG686421) with three major open reading frames (ORFs) and a fourth potential ORF overlapping with the end of ORF3 (Olszewski and Lockhart 2011; Bhat et al. 2016; Rumbou et al. 2018).

Cherry leaf roll virus Taxonomic position Genus: Nepovirus

(CLRV)

Family: Secoviridae

CLRV infection in plants of Betula spp. was reported from Northern Norway, Finland, Eastern Germany, Sweden, and the UK (Cooper and Atkinson 1975; Cooper and Massalski 1984; Jones et al. 1990; Werner et al. 1997; Jalkanen et al. 2007; von Bargen et al. 2009, 2013). The virus-infected birch

Betula spp. (Birch)

277

plants exhibit symptoms of foliar mottle, pale yellow spots, bright green veinbanding, leaf roll, chlorosis, and subsequent necrosis of leaves (Gruntzig et al. 1996). The virus is transmitted by nematode vectors in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants. The virus is seed-transmitted in Betula spp. (Cooper 1976; Cooper et al. 1984). For more details of CLRV, refer to Prunus avium.

Tomato bushy stunt virus Taxonomic position Genus: Tombusvirus

(TBSV)

Family: Tombusviridae

TBSV infection in plants of Betula pendula was reported by Novak and Lanzova (1980). The virus is transmitted by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants. There is no known vector for this virus. For more details of TBSV, refer to Solanum lycopersicum.

References Bhat AI, Hohn T, Selvarajan R (2016) Badnaviruses: the current global scenario. Viruses 8(6):177 Cooper JI (1976) The possible epidemiological significance of pollen and seed transmission in the Cherry leaf roll virus (Betula spp.) complex. Mitt Biol Bundst Ld-Forstw 170:17–22 Cooper JI, Atkinson MA (1975) Cherry leaf roll virus causing a disease of Betula spp. in the United Kingdom. Forestry 48:193–203 Cooper JI, Massalski PR (1984) Viruses and virus-like diseases affecting Betula spp. Proc R Soc Edinb 85B:183–195 Cooper JI, Massalski PR, Edwards M-L (1984) Cherry leaf roll virus in the female gametophyte and seed of birch and its relevance to vertical virus transmission. Ann Appl Biol 105:55–64 Gotlieb AR (1975) Apple mosaic virus infecting yellow birch in Vermont. Proc Am Phytopathol Soc 2:97 Gotlieb AR, Berbee JG (1973) Line pattern of birch caused by Apple mosaic virus. Phytopathology 63:1470–1477 Gruntzig M, Fuchs E, Hentsch T (1996) Occurrence and serological detection of Cherry leaf roll nepovirus (CLRV) and Apple mosaic ilarvirus (ApMV) in Betula spp. J Plant Dis Protect 103:571–581 Jalkanen R, Buttner C, von Bargen S (2007) Cherry leaf roll virus abundant on Betula pubescens in Finland. Silva Fenn 41(4):755–762 Jones AT, Koenig R, Lesemann D-E, Hamacher J, Nienhaus F, Winter S (1990) Serological comparison of isolates of Cherry leaf roll virus from diseased beech and birch trees in a forest decline area in Germany with other isolates of the virus. J Phytopathol 129:339–344 Novak JB, Lanzova J (1980) Demonstration of Tomato bushy stunt virus in some forest tree species and plants. Lesnictví 26:1009–1016 Olszewski NE, Lockhart B (2011) Badnavirus. Caulimoviridae. The Springer index of viruses. Springer, New York, pp 263–269. https://doi.org/10.1007/978-0-387-95919-1_40 Polak Z, Prochazkova Z (1996) Apple mosaic virus associated with decline of silver birch. Ochrana Rostlin – UZPI 32:15–18 Rumbou A, Candresse T, Marais A, Theil S, Langer J, Jalkanen R, Buttner C (2018) A novel Badnavirus discovered from Betula sp. affected by birch leaf-roll disease. PLoS ONE 13(3):e0193888. https://doi.org/10.1371/journal. pone.0193888 von Bargen S, Grubits E, Jalkanen R, Buttner C (2009) Cherry leaf roll virus – an emerging virus in Finland? Silva Fenn 43:727–738 von Bargen S, Dierker L, Rumbou A, Jalkanen R, Buttner C (2013) Sequence diversity of cherry leaf roll virus makes a difference in infected birches in Finland. PPPHE, Satellite Meeting of the IUFRO unit 07.02.04, 29. May 2013, Berlin Werner R, Muhlbach H-P, Buttner C (1997) Detection of Cherry leaf roll nepovirus (CLRV) in birch, beech and petunia by immunocapture RT-PCR using a conserved primer pair. Eur J For Pathol 27:309–318

B

278

Bidens spp. (Beggarticks)

Bidens spp. (Beggarticks) Family: Asteraceae

Bidens mosaic virus Taxonomic position Genus: Potyvirus

Weed Host

(BiMV)

Family: Potyviridae

Geographical distribution BiMV infection in plants of Bidens spp. was first reported from Brazil by Kitajima et al. (1961). The virus spreads in Brazil (Inoue-Nagata et al. 2006). Symptoms and host(s) The virus-infected Bidens pilosa plants exhibit mosaic and necrosis symptoms. Transmission The virus is transmitted by aphid vectors, Myzus persicae, Aphis coreopsidis, and Dactynotus spp. in a non-persistent manner. The virus is mechanically sap-transmissible to several family members producing systemic symptoms and also induces local lesions on Chenopodium spp. No seed transmission is recorded. Virion properties and genome The virions are non-enveloped, flexuous filaments with a modal length of 730 nm and width of 15 nm. The genome consists of a single molecule of linear, positive-sense ssRNA of 9557 nt (KF649336 = NC_023014) encoding a predicted polyprotein of 348 kDa (Sanches et al. 2014; Revers and Garcia 2015; Wylie et al. 2017).

Bidens mottle virus Taxonomic position Genus: Potyvirus

(BiMoV)

Family: Potyviridae

Geographical distribution BiMoV infection in plants of Bidens spp. was reported from Florida and New York (Christie et al. 1968; Youssef et al. 2008). Symptoms and host(s) The virus-infected beggar-ticks plants exhibit mild chlorotic mottling symptoms. The virus naturally infects lettuce, endive, blue lupin, and the foliage plant Fittonia. Transmission The virus is transmitted by aphid vectors, Acyrthosiphon pisum, Aphis craccivora, Aphis spiraecola, Hyadaphis pseudobrassicae, and Myzus persicae in a non-persistent manner. The virus is also mechanically sap-transmissible and not transmitted by seed.

Bidens spp. (Beggarticks)

279

Virion properties and genome The virions contains a single non-enveloped, flexuous filaments 680–900 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9741 nt (AF538686 = NC_014325, EU250210) (Youssef et al. 2008; Revers and Garcia 2015; Wylie et al. 2017).

B Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Bidens pilosa was reported from Australia (Sharman et al. 2015). The virusinfected beggar-ticks leaves exhibit reddening and leaf deformation symptoms. The virus is transmitted by the thrips vectors, is present in/on pollen, and enters into the host through the injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Bidens spp. was reported from Maui Island in Hawaii and China (Cho et al. 1986; Huang et al. 2016). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Cho JJ, Mau RFL, Gonsalves D, Mitchell WC (1986) Reservoir weed hosts of Tomato spotted wilt virus. Plant Dis 70:1014–1017 Christie SR, Edwardson JR, Zettler FW (1968) Characterization and electron microscopy of virus isolated from Bidens and Lepidium. Plant Dis Reptr 52:763–767 Huang CJ, Liu Y, Yu HQ, Liu BZ, Qing L (2016) Bidens pilosa is a natural host of tomato spotted wilt virus in Yunnan Province, China. Plant Dis 100:1957 Inoue-Nagata AK, Oliveira PA, Dutra LS, Nagata T (2006) Bidens mosaic virus is a member of the Potato virus Y species. Virus Genes 33(1):45–49 Kitajima EW, Carvalho AMB, Costa AS (1961) Morfologia do vírus do mosaico do picao. Bragantia, Campinas 20:503–512 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Sanches MM, DeMarchi BR, Spadotti DM, Nozaki DN, Pavan MA, Krause-Sakate R (2014) Biological and molecular characterisation of Bidens mosaic virus supports its assignment as a member of a distinct species in the genus Potyvirus. Arch Virol 159:2181–2183 Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354 Youssef F, Marais A, Candresse T (2008) Partial genome sequence of Bidens mottle virus sheds light on its taxonomy. Arch Virol 153:227–228

280

Bituminaria bituminosa (Arabian pea)

Bituminaria bituminosa (Arabian pea) Family: Fabaceae

Forage crop

Alfalfa mosaic virus

(AMV)

Taxonomic position Genus: Alfamovirus

Family: Bromoviridae

AMV infection in plants of Bituminaria bituminosa was reported from Australia (Jones et al. 2012). The virus-infected Arabian pea plants exhibit bright yellow mosaic symptoms. The virus is transmitted by aphids, such as Acyrthosiphon pisum and Myzus persicae, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Passiflora chlorosis virus Genus: Potyvirus

(PaChV)

Family: Potyviridae

PaChV infection in plants of Bituminaria bituminosa was reported from France, Italy, and Spain (Cardin and Moury 2009). The virus-infected plants exhibit symptoms of mosaic and chlorotic ringspot symptoms on leaves. The virus was transmissible by the green peach aphid (Myzus persicae) to healthy B. bituminosa seedlings in a non-persistent manner. The virus is also mechanically sap-transmissible to a number of hosts. For more details of PaChV, refer to Passiflora spp.

References Cardin L, Moury B (2009) First report of Passiflora chlorosis virus in Bituminaria bituminosa in Europe. Plant Dis 93:196 Jones RAC, Real D, Vincent SJ, Gajda BE, Coutts BA (2012) First report of Alfalfa mosaic virus infecting tedera (Bituminaria bituminosa (L.) C.H. Stirton var. albomarginata and crassiuscula) in Australia. Plant Dis 96:1384

Blainvillea rhomboidea (Blainvillea) Family: Asteraceae

Weed host

Blainvillea yellow spot virus Taxonomic position Genus: Begomovirus

(BlYSV)

Family: Geminiviridae

Geographical distribution BlYSV infection in plants of B. rhomboidea was reported from Brazil (Castillo-Urquiza et al. 2008; Rocha et al. 2013).

Blechum pyramidatum (Green shrimp plant; Browne’s Blechum)

281

Symptoms and host(s) The virus-infected plants exhibit foliar yellow mosaic on the leaves. The virus was also isolated from another weed Physalis spp. Transmission Insect transmission of BlYSV has not been demonstrated. However, in common with all other begomoviruses, BlYSV is likely transmitted by the whitefly vector of begomoviruses, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of BlYSV has not been investigated. In common with all geminiviruses the virions of BlYSV will likely be geminate (twinned icosahedra). The genome of BlYSV is bipartite consisting of two circular, single-stranded DNA components. DNA-A contains 2666 nt (EU710756 = NC_010837) and DNA-B of 2634 nt (EU710757 = NC_010838). Typical of all bipartite begomoviruses native to the New World, the DNA-A component of BlYSVencodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation (Briddon 2001; Castillo-Urquiza et al. 2008; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the New World the DNA-A component of BlYSV encodes five genes, one in the virion-sense and four in the complementary-sense whereas the DNA-B encodes one gene in each orientation (Castillo-Urquiza et al. 2008). The expression and function of these genes has not been investigated for BlYSV.

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Castillo-Urquiza GP, Beserra JE Jr, Bruckner FP, Lima AT, Varsani A, Alfenas-Zerbini P, Murilo Zerbini F (2008) Six novel begomoviruses infecting tomato and associated weeds in Southeastern Brazil. Arch Virol 153(10):1985–1989 Rocha CS, Castillo-Urquiza GP, Lima ATM, Silva FN, Xavier CAD, Hora-Junior BT, Beserra-Junior JEA, Malta AWO, Martin DP, Varsani A, Alfenas-Zerbini P, Mizubuti ESG, Zerbini FM (2013) Brazilian begomovirus populations are highly recombinant, rapidly evolving, and segregated based on geographical location. J Virol 87:5784–5799 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

Blechum pyramidatum (Green shrimp plant; Browne’s Blechum) Family: Acanthaceae

Medicinal

Blechum interveinal chlorosis virus Taxonomic position Genus: Begomovirus

(BleICV)

Family: Geminiviridae

B

282

Blechum pyramidatum (Green shrimp plant; Browne’s Blechum)

Geographical distribution BleICV infection in plants of Blechum pyramidatum was reported from Mexico (Mauricio-Castillo et al. Unpublished - NC_019035). Symptoms and host(s) The virus-infected Browne’s Blechum plants exhibit interveinal chlorosis symptoms. Transmission The transmission of BleICV has not been investigated. It is likely that, in common with other begomoviruses, BleICV will be transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Virion properties and genome The structure of BleICV particles has not been investigated. In common with all geminiviruses the virions of BleICV will likely be geminate (twinned icosahedra). BleICV is a typical New World begomovirus with a bipartite genome consisting of two circular, single stranded DNA components. DNA-A contains 2645 nt (JX827487 = NC_019035) and DNA-B of 2640 nt (JX827488 = NC_019036). Typical of all bipartite begomoviruses native to the New World the DNA A component of BleICV encodes five genes, one in the virion-sense and four in the complementary-sense whereas the DNA B encodes one gene in each orientation (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Blechum yellow vein virus Taxonomic position Genus: Begomovirus

(BleYVV)

Family: Geminiviridae

Geographical distribution BleYVV infection in plants of Blechum pyramidatum was reported from the Philippines (Tsai et al. 2014). Symptoms and host(s) The virus-infected Browne’s Blechum plants exhibit symptoms of yellow/chlorotic leaf veins and shortened internodes. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22 × 38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2764 nt (KF446659) (Tsai et al. 2014; Brown et al. 2015; Zerbini et al. 2017).

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619

Boehmeria nivea (Ramie)

283

Tsai WS, Shih SL, Lee LM, Dolores LM, Kenyon L (2014) First report of a novel Begomovirus associated with yellow vein disease of Browne’s Blechum (Blechum pyramidatum). Plant Dis 98(5):701 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

Blephilia hirsuta (Hairy wood mint) Family: Lamiaceae

Medicinal

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Blephilia hirsuta was reported from North America (Poudel et al. 2010). The virus-infected plants exhibit mottling symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

References Poudel B, Laney AG, Tzanetakis IE (2010) First report of Cucumber mosaic virus infecting Blephilia hirsuta in North America. Plant Dis 94:1070

Boehmeria nivea (Ramie) Family: Urticaceae

Fiber crop

Corchorus golden mosaic virus Taxonomic position Genus: Begomovirus

(CoGMV)

Family: Geminiviridae

CoGMV infection in plants of Boehmeria nivea was reported from Barrackpore (India) (Biswas et al. 2016). The virus-infected plants exhibit symptoms of vein-clearing which was prominent in young leaves, whereas the older leaves show complete yellowing. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of CoGMV, refer to Corchorus spp.

B

284

Boehmeria nivea (Ramie)

Ramie mosaic Yunnan virus Taxonomic position Genus: Begomovirus

(RMYnV)

Family: Geminiviridae

Geographical distribution RMYnV infection in plants of Boehmeria was reported from China (Zhao et al. unpublished KU522485)

Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner.

Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2759 nt (KU522485 = NC_030699) (Briddon 2001; Zerbini et al. 2017).

Tomato leaf curl Hsinchu virus

(ToLCHsV)

Synonyms Ramie mosaic virus (RamMV)

Taxonomic position Genus: Begomovirus

Family: Geminiviridae

RamMV infection in plants of Boehmeria nivea was reported from China (Li et al. 2010). The virusinfected plants exhibit yellow mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ToLCHsV, refer to Solanum lycopersicum.

References Biswas C, Dey MP, Selvarajan R, Bera A, Mitra S, Satpathy S (2016) First report of Corchorus golden mosaic virus (CoGMV) infecting ramie (Boehmeria nivea) in India. Plant Dis 100:541 Briddon RW (2001) Begomovirus. Geminiviridae. The Springer index of viruses. Springer, New York, pp 340–348, https://doi.org/10.1007/3-540-31042-8_55 Li J, Zhang XY, Qian YJ (2010) Molecular characterization of Ramie mosaic virus isolates detected in Jiangsu and Zhejiang provinces, China. Acta Virol 54(3):225–228 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

Boerhavia erecta (Erect spiderling)

285

Boerhavia erecta (Erect spiderling) Family: Nyctaginaceae

Medicinal

B Boerhavia yellow spot virus Taxonomic position Genus: Begomovirus

(BoYSV)

Family: Geminiviridae

Geographical distribution BoYSV infection in plants of Boerhavia erecta was reported from the Yucatan Peninsula, Mexico (Hernandez-Zepeda et al. 2007). Symptoms and host(s) The virus-infected plants exhibit foliar chlorotic spots and leaf crumpling symptoms. The host range of the virus has not been investigated. Transmission The transmission of BoYSV has not been investigated. It is likely that, in common with other begomoviruses, BoYSV will be transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. The virus is not mechanically sap-transmissible. Virion properties and genome The structure of BoYSV particles has not been investigated. In common with all geminiviruses the virions of BoYSV will likely be geminate (twinned icosahedra). The genome of BoYSV is not well characterized yet. Only the sequence of a single circular component, homologous to the DNA-A genome (2621 nt, EF121755) of monopartite begomoviruses has been determined. The majority of begomoviruses native to the New World are bipartite, suggesting that the sequence of the genome of BoYSV is incomplete – missing the DNA-B component. The only available sequence of BoYSV encodes the five genes typically encoded by the DNA-A components of bipartite begomoviruses or the genomes of monopartite begomoviruses native to the New World (Briddon 2001; Hernandez-Zepeda et al. 2007; Brown et al. 2015; Zerbini et al. 2017).

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Hernandez-Zepeda C, Idris AM, Carnevali G, Brown JK, Moreno-Valenzuela OA (2007) Preliminary identification and coat protein gene phylogenetic relationships of begomoviruses associated with native flora and cultivated plants from the Yucatan Peninsula of Mexico. Virus Genes 35:825–833 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

286

Borago officinalis (Borage, Starflower)

Borago officinalis (Borage, Starflower) Family: Boraginaceae

Medicinal

Alfalfa mosaic virus

(AMV)

Taxonomic position Genus: Alfamovirus

Family: Bromoviridae

AMV infection in plants of Borago officinalis was reported from Spain (Mallor et al. 2002). The virusinfected starflower plants exhibit foliar yellow mosaic symptoms. The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Clover yellow vein virus Taxonomic position Genus: Potyvirus

(ClYVV)

Family: Potyviridae

ClYVV infection in plants of Borago officinalis was reported from Spain (Luis-Arteaga et al. 1996). The virus-infected starflower plants exhibit symptoms of mosaic and deformation of the leaves. The virus is transmitted by an aphid vector, Myzus persicae, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ClYVV, refer to Trifolium spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Borago officinalis was reported from Spain (Luis-Arteaga et al. 1988). The virus-infected starflower plants exhibit stunting of the plant, leaf deformation, and mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

References Luis-Arteaga M, Rodriguez-Cerezo E, Maestro C, Garcia-Arenal F (1988) Detection and characterization of an isolate of Cucumber mosaic virus (CMV) infecting borage (Borago officinalis L.) in Spain. Plant Dis 72:265–267 Luis-Arteaga M, Garcia-Arenal F, Rodriguez-Cerezo E (1996) Characterization of a strain of Clover yellow vein potyvirus infecting borage (Borago officinalis L.) in Spain. Plant Pathol 45:38–44 Mallor C, Luis-Arteaga M, Cambra MA, Fernandez-Cavada S (2002) Natural infection of field-grown borage (Borago officinalis) by Alfalfa mosaic virus in Spain. Plant Dis 86(6):698

Bougainvillea spp.

287

Bougainvillea spp. Family: Nyctaginaceae

Ornamental

B Bougainvillea chlorotic veinbanding virus

(BCVBV)

Synonyms Bougainvillea bacilliform virus; Bougainvillea chlorotic spot virus; Bougainvillea spectabilis chlorotic veinbanding virus Taxonomic position Genus: Badnavirus

Family: Caulimoviridae

Geographical distribution BCVBV infection in plants of Bougainvillea spectabilis was first reported from Brazil (Chagas et al. 2001). The virus spreads in India, Brazil, and Taiwan (Yamashita et al. 2004; Rivas et al. 2005; Tsai et al. 2005, 2008; Baranwal et al. 2010). Symptoms and host(s) The virus-infected plants develop symptoms such as mottling, chlorosis, veinbanding, severe leaf distortion, and stunting. Transmission The virus is not mechanically sap-transmissible, but transmitted through grafting. Virion properties and genome The virions are bacilliform, non-enveloped, about 28 nm in diameter, and varying in length between 130 and 150 nm. The genome consists of a single molecule of non-covalently closed circular doublestranded DNA of 8759 bp (EU034539 = NC_011592) (Olszewski and Lockhart 2011; Bhat et al. 2016).

Clerodendron yellow mosaic virus Taxonomic position Genus: Begomovirus

(ClYMV)

Family: Geminiviridae

ClYMV infected plants of Bougainvillea peruviana were reported from India (Nehra et al. 2014). The virus-infected plants exhibit symptoms of leaf curling and stunted growth. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ClYMV, refer to Clerodendrum spp.

288

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Bougainvillea spp.

(CMV)

Family: Bromoviridae

CMV infection in plants of Bougainvillea spectabilis was reported from Iran (Shahmohammadi et al. 2015). The virus-infected plants exhibit mosaic symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Bougainvillea spectabilis was reported from Iran (Shahraeen et al. 2002; Ghotbi 2013). The virus-infected plants exhibit necrotic spots, leaf yellowing, ringspots, necrotic veinclearing, wilting, and dwarf symptoms. The virus is transmitted by thrips vectors in a persistentpropagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato yellow ring virus

(TYRV)

Synonyms Tomato Varamin virus (ToVV) Taxonomic position TYRV is a tentative member of the genus Orthotospovirus and family Tospoviridae. ToVV was reported infecting plants of Bougainvillea spp. in Iran (Ghotbi et al. 2005). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of TYRV, refer to Solanum lycopersicum.

References Baranwal VK, Arya M, Singh J (2010) First report of two distinct badnaviruses associated with Bougainvillea spectabilis in India. J Gen Plant Pathol 76:236–239 Bhat AI, Hohn T, Selvarajan R (2016) Badnaviruses: the current global scenario. Viruses 8(6):177 Chagas CM, Alexandre MAV, Duarte LML, Rivas EB, Tombolato AFC (2001) Badnavirus-like particles associated with chlorotic vein-banding symptoms in Bougainvillea spectabilis. Virus Rev Res 6:153 Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iran J Plant Pathol 49:41 Ghotbi T, Sharaeen N, Winter S (2005) Occurrence of tospoviruses in ornamental and weed species in Markazi and Tehran provinces in Iran. Plant Dis 89:425–429 Nehra C, Sahu AK, Marwal A, Gaur RK (2014) Natural occurrence of Clerodendron yellow mosaic virus on Bougainvillea in India. New Dis Rep 30:19

Bouvardia spp.

289

Olszewski NE, Lockhart B (2011) Badnavirus. Caulimoviridae. In: The Springer index of viruses. Springer, New York, pp 263–269. https://doi.org/10.1007/978-0-387-95919-1_40 Rivas EB, Duarte LM, Alexandre MAV, Fernandes FM, Harakava R, Chagas CM (2005) A new Badnavirus species detected in Bougainvillea in Brazil. J Gen Plant Pathol 71(6):438–440 Shahmohammadi N, Dizadji A, Habibi MK, Nateqi M (2015) First report of Cucumber mosaic virus infecting Bougainvillea spectabilis, Coleus blumei, Kalanchoe blossfeldiana and Zinnia elegans in Iran. J Plant Pathol 97(2):394 Shahraeen N, Ghotbi T, Mehraban AH (2002) Occurrence of Impatiens necrotic spot virus in ornamentals in Mahallat and Tehran provinces in Iran. Plant Dis 86:694 Tsai C-H, Su H-J, Liao Y-C, Hung T-H (2005) First report of Bougainvillea spectabilis chlorotic vein-banding virus infecting bougainvillea plants in Taiwan. Plant Dis 89:1363 Tsai C-H, Su H-J, Wu M-L, Feng Y-C, Hung T-H (2008) Identification and detection of Bougainvillea spectabilis chlorotic vein-banding virus in different bougainvillea cultivars in Taiwan. Ann Appl Biol 153:187–193 Yamashita S, Ferreira PTO, Figueiredo DV, Brioso PST, Kitajima EW (2004) Occurrence of a badnavirus in Bougainvillea in Brazil. Summa Phytopathol 30:68–68

Bouvardia spp. Family: Rubiaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in flowers of pink- or red-flowered Bouvardia domestica plants was reported from the Netherlands (Hakkaart 1979). The virus-infected plants show dull or faded flower color, without any visible foliar symptoms; red-flowered plants did not always show floral symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sapinoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Tomato chlorotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(TCSV)

Family: Tospoviridae

TCSV infection in plants of Bouvardia spp. was reported from Brazil (Rivas et al. 2002). The virusinfected plants exhibit mild mosaic symptoms. The virus is transmitted by thrips vectors in a persistentpropagative manner, and also by mechanical sap-inoculation. For more details of TCSV, refer to Solanum lycopersicum.

References Hakkaart FA (1979) Virus in bouvardias can be controlled by testing and selection. Vakbl Bloemisterij 34(32):40–41 Rivas EB, Galleti SR, Alexandre MAV, Duarte LML, Seabra PV, Estelita MEM (2002) Tomato chlorotic spot virus (Tospovirus) in dieffenbachia and bouvardia. Virus Rev Res 7:22–30

B

290

Brachiaria spp.

Brachiaria spp. Family: Poaceae

Forage crop

Johnsongrass mosaic virus

(JGMV)

Synonyms Guinea grass mosaic virus (GGMV) Taxonomic position Genus: Potyvirus

Family: Potyviridae

GGMV infection in plants of Brachiaria spp. was reported from Colombia and Brazil (Morales et al. 1994). The virus-infected plants of Brachiaria brizantha exhibit symptoms of stippling, chlorotic streaks on leaves, and characteristic eye-shaped lesions and develop variegation in the later stages of an infection. The virus is transmitted by aphid vectors, Hysteroneura setariae and Rhopalosiphum maidis, in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not seed-borne in Brachiaria spp. For more details of JGMV, refer to Sorghum halepense.

Rice necrosis mosaic virus Taxonomic position Genus: Bymovirus

(RNMV)

Family: Potyviridae

RNMV infection in plants of Brachiaria ramosa was reported from India (Ghosh 1981). The virus is transmitted by a fungal vector, in a persistent manner. The virus is also transmitted by mechanical sap-inoculation. For more details of RNMV, refer to Oryza sativa.

Sugarcane mosaic virus Taxonomic position Genus: Potyvirus

(SCMV)

Family: Potyviridae

SCMV infection in plants of Brachiaria piligera was reported from Queensland (Australia) (Srisink et al. 1993). The virus-infected plants exhibit mosaic symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SCMV, refer to Saccharum officinarum.

References Ghosh SK (1981) Weed hosts of rice necrosis mosaic virus. Plant Dis 65:602–603 Morales FJ, Castano M, Velasco AC, Arroyave J (1994) Detection of a Potyvirus related to Guinea grass mosaic virus infecting Brachiaria spp. in South America. Plant Dis 78:425–428 Srisink S, Noon DF, Teakle DS, Ryan CC (1993) Brachiaria piligera and Sorghum verticilliflorum are natural hosts of two different strains of sugarcane mosaic virus in Australia. Australas Plant Pathol 22:94–97

Brassica carinata (Ethiopian Mustard)

291

Brassica campestris var. purpuraria (Purple Cai-tai) Family: Brassicaceae

Vegetable

B Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Brassica campestris var. purpuraria was reported from China (Cai et al. 2017). The virus-infected purple cai-tai plants show mosaic and shrinking on leaves and dwarfing symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

References Cai L, Zhang YC, Chen YS, Hong N, Xu WX (2017) First report of cucumber mosaic virus infecting Brassica campestris var. purpuraria in China. Plant Dis 101:1684

Brassica carinata (Ethiopian Mustard) Family: Brassicaceae

Turnip mosaic virus Taxonomic position Genus: Potyvirus

Oilseed

(TuMV)

Family: Potyviridae

TuMV infection in plants of Brassica carinata was reported from the USA (Babu et al. 2013). The virus-infected plants exhibit mosaic, ringspot, mottling, and puckering symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

References Babu B, Dankers H, George S, Wright D, Marois J, Paret M (2013) First report of Turnip mosaic virus infecting Brassica carinata (Ethiopian mustard) in the United States. Plant Dis 97:1664

292

Brassica juncea (Indian mustard/Rai)

Brassica juncea (Indian mustard/Rai) Family: Brassicaceae

Oil seed

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Brassica juncea was reported from India and Taiwan (Green and Deng 1985; Haq et al. 1994; Singh et al. 2015). The virus-infected Indian mustard plants exhibit systemic mosaic and leaf deformation symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

Turnip yellows virus Taxonomic position Genus: Polerovirus

(TuYV)

Family: Luteoviridae

TuYV infection in plants of Brassica juncea was reported from Rajasthan (India) and China (Xiang et al. 2009, 2011; Sharma et al. 2013). The virus-infected Indian mustard plants exhibit symptoms of yellow mosaic and reddening of leaves. The virus is transmitted by a number of aphid species in a circulative and non-propagative manner. The virus is not mechanically sap-transmissible. For more details of TuYV, refer to Brassica rapa.

Turnip yellow mosaic virus

(TYMV)

Synonyms Rai mosaic virus Taxonomic position Genus: Tymovirus

Family: Tymoviridae

TYMV infection in plants of Brassica juncea was reported from West Bengal (India) (Biswas and Chowdhury 2005). The virus-infected Indian mustard plants exhibit systemic yellow to light green mosaic symptoms. The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sap-inoculation, and by grafting. For more details of TYMV, refer to Brassica rapa.

References Biswas KK, Chowdhury A (2005) Rai mosaic disease in rai (Brassica juncea) and radish (Raphanus sativus) in the Himalayan hilly regions of West Bengal. J Mycopathol Res 43:263–266

Brassica napus (Canola/Oilseed rape/Rapeseed)

293

Green SK, Deng TC (1985) Turnip mosaic virus strains in cruciferous hosts in Taiwan. Plant Dis 69:28–31 Haq QMR, Srivastava KM, Raizada RK, Singh BP, Jain RK, Mishra A, Shukla DD (1994) Biological, serological and coat protein properties of a strain of Turnip mosaic virus causing a mosaic disease of Brassica campestris and B. juncea in India. J Phytopathol 140:55–64 Sharma P, Verma RK, Mishra R, Choudhary DK, Gaur RK (2013) First report of Turnip yellow virus (TuYV) in Brassica juncea (Indian mustard) in India. New Dis Rep 27:21 Singh R, Banerjee A, Sharma SK, Bhagawati R, Baruah S, Ngachan SV (2015) First report of Turnip mosaic virus occurrence in cole crops (Brassica spp) from Arunachal Pradesh, India. Virus Dis 26:211–213 Xiang H, Shang Q, Han C, Li D, Yu J (2009) Detection of Turnip yellows virus in eight cruciferous crops in Mainland China. Phytopathology 99:S144 Xiang H-Y, Dong S-W, Shang Q-X, Zhou C-J, Wei Li D, Yu J-L, Han C-G (2011) Molecular characterization of two genotypes of a new Polerovirus infecting brassicas in China. Arch Virol 156:2251–2255

Brassica napus (Canola/Oilseed rape/Rapeseed) Family: Brassicaceae

Oil seed

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

BWYV was reported in plants of Brassica napus from south-west Australia, the UK, and Iran (Smith and Hinckes 1985; Walsh and Tomlinson 1985; Walsh 1986; Hill et al. 1989; Hardwick et al. 1994; Coutts and Jones 2000; Jones and Hawkes 2002; Shahraeen et al. 2003; Tabarestani et al. 2010; Shahraeen 2012). The virus-infected canola plants show interveinal chlorosis on lower and intermediate leaves. Leaves are frequently purple and plants ripen prematurely. Infection with this virus in canola can cause partial plant dwarfing and reddening of lower leaves, but infected plants are usually devoid of obvious symptoms (Jay et al. 1995). The virus is transmitted by the aphids Myzus persicae and Macrosiphum euphorbiae and possibly by other aphid vectors in a persistent manner. No mechanical sap-transmission and also no seed transmission. For more details of BWYV, refer to Beta vulgaris.

Broccoli necrotic yellows cytorhabdovirus Taxonomic position Genus: Cytorhabdovirus

(BNYV)

Family: Rhabdoviridae

BNYV infection in plants of Brassica napus was reported from the UK (Walsh 1986; Walsh and Tomlinson 1985). The virus is transmitted by aphid vectors in a persistent, propagative manner, and also by mechanical sap-inoculation. For more details of BNYV, refer to Brassica oleracea var. italica.

Cauliflower mosaic virus Taxonomic position Genus: Caulimovirus

(CaMV)

Family: Caulimoviridae

B

294

Brassica napus (Canola/Oilseed rape/Rapeseed)

CaMV was reported on Brassica napus plants from Bulgaria, New Zealand, Iran, south-west Australia, the UK, and the USA (Kolte 1985; Walsh and Tomlinson 1985; Walsh 1986; Hardwick et al. 1994; Coutts and Jones 2000; Shahraeen et al. 2003; Farzadfar et al. 2005; Sahandi et al. 2005; Ghaderi et al. 2009a, b; Tabarestani et al. 2010; Shahraeen 2012; Ebrahim-Ghomi 2014). The virus-infected canola plants exhibit chlorotic ringspots and mottling symptoms. The virus is transmitted by about 27 aphid species such as Myzus persicae and Brevicoryne brassicae in semi-persistent manner. The virus is also mechanically sap-transmissible with difficulty, and cruciferous plants are susceptible. For more details of CaMV, refer to Brassica oleracea var. botrytis.

Citrus exocortis viroid Taxonomic position Genus: Pospiviroid

(CEVd)

Family: Pospiviroidae

CEVd infection in plants of Brassica napus was reported from Spain (Fagoaga and Duran-Vila 1996). The viroid is mechanically sap-transmissible and also through contaminated tools. For more details of CEVd, refer to Citrus spp.

Croton yellow vein mosaic virus Taxonomic position Genus: Begomovirus

(CroYVMV)

Family: Geminiviridae

CroYVMV infection in plants of Brassica napus was reported from India (Roy et al. 2013). The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. For more details of CroYVMV, refer to Codiaeum variegatum.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Brassica napus was reported from Iran (Shahraeen et al. 2003). The virusinfected canola plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a nonpersistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Brassica napus was reported from Iran (Shahraeen 2012; Shahraeen et al. 2003). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by

Brassica napus (Canola/Oilseed rape/Rapeseed)

295

mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Brassica napus is widespread in distribution (Shen and Pu 1965; Rao et al. 1977; Walsh and Tomlinson 1985; Walsh 1986; Hardwick et al. 1994; Coutts and Jones 2000; Shahraeen et al. 2003; Ghorbani et al. 2006; Tabarestani et al. 2010; Sabokkhiz et al. 2012; Shahraeen 2012). The virus-infected canola plants exhibit mosaic and stunting symptoms. The virus is transmitted by aphid vectors, Myzus persicae, Brevicoryne brassicae, Lipaphis erysimi, Aphis craccivora, and Aphis gossypii, in a non-persistent manner and being retained by aphids for not more than 4 h after acquisition and usually much less. The virus is mechanically sap-transmissible to a large number of test plants belonging to more than nine families. For more details of TuMV, refer to Brassica rapa.

Turnip yellow mosaic virus Taxonomic position Genus: Tymovirus

(TYMV)

Family: Tymoviridae

TYMV infection in plants of Brassica napus was reported from the Czech Republic (Spak et al. 1993). The virus-infected canola plants exhibit yellow mosaic symptoms. The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sap-inoculation, and by grafting. The virus is seed-transmitted up to 1.5% in different cultivars of rapeseed (Spak et al. 1993). For more details of TYMV, refer to Brassica rapa.

Turnip yellows virus Taxonomic position Genus: Polerovirus

(TuYV)

Family: Luteoviridae

TuYV infection in plants of Brassica napus was reported from the UK, Austria, China, Germany, and Serbia (Graichen and Schliephake 1999; Graichen et al. 2000; Xiang et al. 2009, 2011; Milosevic et al. 2015, 2016). The virus-infected canola plants exhibit symptoms of leaf yellowing and reddening of margins followed by leaf distortion. The virus is transmitted by a number of aphid species in a circulative and non-propagative manner (Schliephake et al. 2000). The virus is not mechanically saptransmissible. For more details of TuYV, refer to Brassica rapa.

Youcai mosaic virus Synonyms Oilseed rape mosaic virus

(YoMV)

B

296

Taxonomic position Genus: Tobamovirus

Brassica napus (Canola/Oilseed rape/Rapeseed)

Family: Virgaviridae

Geographical distribution YoMV infection in plants of Brassica napus was reported from China (Cai et al. 2009). Symptoms and host(s) The virus-infected canola plants exhibit mosaic symptoms. Transmission There is no known vector for this virus. The virus is mechanically sap-transmissible and also through contact between plants. Virion properties and genome The virions are non-enveloped, rigid helical rods with a helical symmetry, about 18 nm in diameter and 300–310 nm in length. The genome is a monopartite, linear, positive-sense ssRNA of 6303 nt (U30944 = NC_004422). The 30 terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G50 pppG) (Aguilar et al. 1996; Zaitlin 2011; Adams et al. 2017).

References Adams MJ, Adkins S, Bragard C, Gilmer D, Li D, MacFarlane SA, Wong S-M, Melcher U, Ratti C, Ryu KH, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Virgaviridae. J Gen Virol 98:1999–2000 Aguilar I, Sanchez F, Martin Martin A, Martinez-Herrera D, Ponz F (1996) Nucleotide sequence of Chinese rape mosaic virus (oilseed rape mosaic virus), a crucifer Tobamovirus infectious on Arabidopsis thaliana. Plant Mol Biol 30(1):191–197 Cai L, Chen K, Zhang X, Yan L, Hou M, Xu Z (2009) Biological and molecular characterization of a crucifer Tobamovirus infecting oilseed rape. Biochem Genet 47:451–461 Coutts BA, Jones RAC (2000) Viruses infecting canola (Brassica napus) in south-west Australia: incidence, distribution, spread, and infection reservoir in wild radish (Raphanus raphinistrum). Aust J Agric Res 51:925–936 Ebrahim-Ghomi M (2014) Study on distribution and detection of Cauliflower mosaic virus (CaMV) in Dezful region of Iran. Int J Biosci 4:271–275 Fagoaga C, Duran-Vila N (1996) Naturally occuring variants of Citrus exocortis viroid in vegetable crops. Plant Pathol 45:45–53 Farzadfar SH, Pourrahim R, Golnaraghi AR, Ahoonmanesh A (2005) Occurrence of Cauliflower mosaic virus in different cruciferous plants in Iran. Plant Pathol 54:810 Ghaderi M, Shahraeen N, Rakhshandehroo F (2009a) Serological and molecular identification of Cauliflower mosaic virus (CaMV) on Canola. 5th Iranian Congress of Virology and 1st Congress of Vaccine. Razi Vaccine and Serum Research Institute. 12–14 May 2009, Karaj-Iran Ghaderi M, Shahraeen N, Rakhshandehroo F (2009b) Identification and study of some properties of Cauliflower mosaic virus isolates infecting canola plants. Biotechnology Congress 7–8 Oct 2009, Karaj-Iran Ghorbani S, Shahraeen N, Dehghanyar H, Sahandi A, Pourrahim R (2006) Serodiagnosis and purification of Turnip mosaic virus isolated from oilseed rape. Iran J Biol 20(1):61–70 Graichen K, Schliephake E (1999) Infestation of winter oilseed rape by Turnip yellows luteovirus and its effect on yield in Germany. In: 10th International Rapeseed Congress – New Horizons from an Old Crop. pp 131 Graichen K, Rabenstein F, Kurtze E (2000) The occurrence of Turnip yellows virus in winter oilseed rape in Austria. Pflanzenschutzberichte 59:35–46 Hardwick NV, Davies JML, Wright DM (1994) The incidence of three virus diseases of winter oilseed rape in England and Wales in the 1991/92 and 1992/93 growing season. Plant Pathol 43:1045–1049 Hill A, Lane A, Hardwick NV (1989) The incidence and importance of Beet western yellows virus in oilseed rape. Asp Appl Biol 23:311–318 Jay CN, Rossall S, Smith HG (1995) The effects of Beet western yellows virus on the growth and yield of oil seed rape (Brassica napus). J Agric Sci 133:131–139

Brassica napus var. napobrassica or Brassica napus sub spp. rapifera (Swedish turnip/Rutabaga)

297

Jones R, Hawkes J (2002) Yield losses caused when Beet western yellows virus infects canola, Agribusiness Crop Updates. Department of Agriculture, Western Australia Kolte SJ (1985) Rapeseed. In: Diseases of annual edible oilseed crops, vol II. CRC Press, Boca Raton, FL, USA, p 135 Milosevic D, Marjanovic-Jeromela A, Ignjatov M, Jovicic D, Stankovic I, Bulajic A, Krstic B (2015) First report of Turnip yellows virus on oilseed rape in Serbia. Plant Dis 99:1869 Milosevic D, Ignjatov M, Nikolic Z, Stankovic I, Bulajic A, Marjanovic-Jeromela A, Krstic B (2016) The presence of Turnip yellows virus in oilseed rape (Brassica napus L.) in Serbia. Pestic Phytomed 31: 37–44 Rao DV, Hiruki C, Chen MH (1977) A mosaic disease of rape in Alberta caused by Turnip mosaic virus. Plant Dis Reptr 61:1074–1076 Roy A, Spoorthi P, Bag MK, Prasad TV, Singh R, Dutta M, Mandal B (2013) A Leaf Curl Disease in Germplasm of Rapeseed-Mustard in India: Molecular Evidence of a Weed-Infecting Begomovirus–Betasatellite Complex Emerging in a New Crop. Journal of Phytopathology 161: 522–535 Sabokkhiz MA, Jafarpour B, Shahriari Ahmadi F, Tarighi S (2012) Identification of Turnip mosaic virus isolated from canola in northeast area of Iran. Afr J Biotech 11:14553–14560 Sahandi A, Shahraeen N, Ghorbani S, Pourrahim R (2005) Identification and studies of some biological and serological properties of Cauliflower mosaic virus (CaMV) isolated from oilseed rape from West Azarbaejan province. J Agric Sci 1(5):101–112 Schliephake E, Graichen K, Rabenstein F (2000) Investigations on the vector transmission of the Beet mild yellowing virus (BMYV) and the Turnip yellows virus (TuYV). J Plant Dis Protect 107:81–87 Shahraeen N (2012) An overview of oilseed rape (canola) virus diseases in Iran. Int Res J Microbiol 3:24–28 Shahraeen N, Farzadfar SH, Lesemann DE (2003) Incidence of viruses infecting winter oilseed rape (Brassica napus ssp. oleifera) in Iran. J Phytopathol 151:614–616 Shen SH, Pu ZG (1965) A preliminary study of the two strain of Turnip mosaic virus on rape in Kiangsu province. Acta Phytophylac SN 4(1):35 Smith GH, Hinckes A (1985) Studies on Beet western yellows virus in oilseed rape (Brassica napus ssp. oleifera) and sugar beet (Beta vulgaris). Ann Appl Biol 107:473–484 Spak J, Kubelkova DA, Hnilicka E (1993) Seed transmission of Turnip yellow mosaic virus in winter turnip and winter oil seed rapes. Ann Appl Biol 123:33–35 Tabarestani AZ, Shamsbakhsh M, Safaei N (2010) Distribution of three important aphid borne canola viruses in Golestan province. Iran J Plant Protect Sci 41:251–259 Walsh JA (1986) Virus diseases of oilseed rape and their control. British Crop Protection Conference. Pests Dis 2:737–743 Walsh JA, Tomlinson JA (1985) Viruses infecting winter oilseed rape (Brassica napus ssp. Oleifera). Ann Appl Biol 107:485–495 Xiang H, Shang Q, Han C, Li D, Yu J (2009) Detection of Turnip yellows virus in eight cruciferous crops in Mainland China. Phytopathology 99:S144 Xiang H-Y, Dong S-W, Shang Q-X, Zhou C-J, Wei Li D, Yu J-L, Han C-G (2011) Molecular characterization of two genotypes of a new polerovirus infecting brassicas in China. Arch Virol 156:2251–2255 Zaitlin M (2011) Tobamovirus. In: Tidona C, Darai G (eds) The Springer Index of Viruses. Springer, New York, NY, https://doi.org/10.1007/978-0-387-95919-1.

Brassica napus var. napobrassica or Brassica napus sub spp. rapifera (Swedish turnip/Rutabaga) Family: Brassicaceae

Turnip mosaic virus Taxonomic position Genus: Potyvirus

Root vegetable

(TuMV)

Family: Potyviridae

TuMV infection in plants of Brassica napus sub spp. rapifera was reported from Canada (Stobbs and Shattuck 1989). The virus-infected rutabaga plants exhibit symptoms of vein chlorosis in young leaves,

B

298

Brassica napus var. silvestris (Winter turnip rape)

secondary symptom of chlorotic, pale yellow spots that later merge into yellow zones on larger portions of the leaf surface. Finally, the green chlorophyll disappears or remains only in traces. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

Turnip yellows virus Taxonomic position Genus: Polerovirus

(TuYV)

Family: Luteoviridae

TuYV infection in plants of Brassica napobrassica was reported from China (Xiang et al. 2011). The virus-infected rutabaga plants exhibit yellowing or leaf roll symptoms. The virus is transmitted by a number of aphid species in a circulative and non-propagative manner. The virus is not mechanically sap-transmissible. For more details of TuYV, refer to Brassica rapa.

References Stobbs LW, Shattuck VI (1989) Turnip mosaic virus strains in southern Ontario, Canada. Plant Dis 73:208–212 Xiang H-Y, Dong S-W, Shang Q-X, Zhou C-J, Wei Li D, Yu J-L, Han C-G (2011) Molecular characterization of two genotypes of a new Polerovirus infecting brassicas in China. Arch Virol 156:2251–2255

Brassica napus var. silvestris (Winter turnip rape) Family: Brassicaceae

Oil seed

Turnip yellow mosaic virus Taxonomic position Genus: Tymovirus

(TYMV)

Family: Tymoviridae

TYMV infection in plants of Brassica napus var. silvestris was reported from the Czech Republic (Spak et al. 1993). The virus-infected winter turnip rape plants exhibit yellow mosaic symptoms. The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sap-inoculation, and by grafting. The virus is seed-transmitted between 1.6% and 8.3% in different cultivars of winter turnip rape (Spak et al. 1993). For more details of TYMV, refer to Brassica rapa.

Turnip yellows virus Taxonomic position Genus: Polerovirus

(TuYV)

Family: Luteoviridae

Brassica nigra (Black mustard)

299

TuYV infection in plants of Brassica napus var. silvestris was reported from Germany (Graichen and Schliephake 1999). The virus-infected winter turnip rape plants exhibit conspicuous anthocyanous discoloration of the leaves and strong reddening symptoms. The virus is transmitted by a number of aphid species in a circulative and non-propagative manner. The virus is not mechanically sap-transmissible. For more details of TuYV, refer to Brassica rapa.

References Graichen K, Schliephake E (1999) Infestation of winter oil seed rape by Turnip yellows luteovirus and its effects on yield in Germany. Proceedings of 10th international rape seed congress, Canberra, Australia Spak J, Kubelkova DA, Hnilicka E (1993) Seed transmission of Turnip yellow mosaic virus in winter turnip and winter oil seed rapes. Ann Appl Biol 123:33–35

Brassica nigra (Black mustard) Family: Brassicaceae

Spice crop

Beet curly top virus Taxonomic position Genus: Curtovirus

(BCTV)

Family: Geminiviridae

BCTV infection in plants of Brassica nigra was reported from California (USA) (Creamer et al. 1996). The virus is transmitted by the leafhopper vector Circulifer tenellus in a persistent (circulative, nonpropagative) manner. The virus is not mechanically sap-transmissible. For more details of BCTV, refer to Beta vulgaris.

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

BWYV infection in plants of Brassica nigra was reported from Dorset (UK) (Thurston et al. 2001). The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmitted by mechanical sap-inoculation. For more details of BWYV, refer to Beta vulgaris.

Cauliflower mosaic virus Taxonomic position Genus: Caulimovirus

(CaMV)

Family: Caulimoviridae

B

300

Brassica nigra (Black mustard)

CaMV infection in plants of Brassica nigra was reported from Dorset (UK) (Thurston et al. 2001). The virus is transmitted by aphid vectors in a semi-persistent manner, and also by mechanical sapinoculation. For more details of CaMV, refer to Brassica oleracea var. botrytis.

Turnip crinkle virus

(TCV)

Taxonomic position Genus: Betacarmovirus

Family: Tombusviridae

TCV infection in plants of Brassica nigra was reported from Dorset (UK) (Thurston et al. 2001). The virus is transmitted by beetle vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TCV, refer to Brassica rapa.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Brassica nigra was reported from Dorset (UK) (Thurston et al. 2001). The virus is transmitted by the aphid vector Myzus persicae in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

Turnip rosette virus Taxonomic position Genus: Sobemovirus

(TRoV)

Family: Solemoviridae

TRoV infection in plants of Brassica nigra was reported from Dorset (UK) (Thurston et al. 2001). The virus is transmitted by beetle vectors and also by mechanical sap-inoculation. For more details of TRoV, refer to Brassica rapa.

Turnip yellow mosaic virus Taxonomic position Genus: Tymovirus

(TYMV)

Family: Tymoviridae

TYMV infection in plants of Brassica nigra was reported from Dorset (UK) (Thurston et al. 2001). The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of TYMV, refer to Brassica rapa.

Brassica oleracea var. acephala (Collard greens)

301

References Creamer R, Lugue-Williams M, Howo M (1996) Epidemiology and incidence of Beet curly top geminivirus in naturally infected weed hosts. Plant Dis 80:533–535 Thurston MI, Pallett DW, Cortina-Borja M, Edwards ML, Raybould AF, Cooper JI (2001) The incidence of viruses in wild Brassica nigra in Dorset (UK). Ann Appl Biol 139:277–284

Brassica oleracea var. acephala (Collard greens) Family: Brassicaceae

Vegetable

Cabbage leaf curl virus Taxonomic position Genus: Begomovirus

(CabLCV)

Family: Geminiviridae

CabLCV was detected in plants of Brassica oleracea var. acephala in Georgia, USA (Mandal et al. 2001). The virus-infected collard green plants show yellow spots, vein-clearing, mosaic, curling, and puckering on leaves. The virus is transmitted by the whitefly vector, Bemisia argentifolii, in a circulative non-propagative manner. The virus is mechanically transmissible to cabbage and collard green plants under experimental conditions. For more details of CabLCV, refer to Brassica oleracea var. capitata.

Cauliflower mosaic virus Taxonomic position Genus: Caulimovirus

(CaMV)

Family: Caulimoviridae

CaMV infection in plants of Brassica oleracea var. acephala was reported from the USA and Iran (Khan and Demski 1982; Farzadfar et al. 2007). The virus-infected collard green plants exhibit vein chlorosis and banding symptoms. The virus is transmitted by aphid vectors such as Brevicoryne brassicae and Myzus persicae in a semi-persistent manner, and also by mechanical sap-inoculation. The virus is not transmitted by seed. For more details of CaMV, refer to Brassica oleracea var. botrytis.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV is known to infect Brassica oleracea var. acephala in the USA, Turkey, and Brazil (Khan and Demski 1982; Sevik 2016). The symptoms of the infected collard green plants include mild mottling, mosaic, and small chlorotic ringspots and blisters. The virus is transmitted by aphid vectors in a non-

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Brassica oleracea var. botrytis (Cauliflower)

persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

References Farzadfar SH, Ahoomanesh A, Mosahebi GH, Pourrahim AR, Golnaraghi AR (2007) Occurrence and distribution of Cauliflower mosaic virus on cruciferous plants in Iran. Plant Pathol J 6:22–29 Khan MA, Demski JW (1982) Identification of turnip mosaic and cauliflower mosaic viruses naturally infecting collards. Plant Dis 66:253–256 Mandal B, Langston DB, Pappu HR, Beard GH, Kucharek TA, Flanders JT, Whiddon JP, Smith JE, Kelley WT (2001) First report of Cabbage leaf curl virus (Family Geminiviridae) in Georgia. Plant Dis 85:561 Sevik MA (2016) Turnip mosaic virus infecting kale plants in Ordu, Turkey. Phyton 85:231–235

Brassica oleracea var. botrytis (Cauliflower) Family: Brassicaceae

Vegetable

Broccoli necrotic yellows cytorhabdovirus Taxonomic position Genus: Cytorhabdovirus

(BNYV)

Family: Rhabdoviridae

BNYV was first reported in plants of Brassica oleracea var. botrytis from England by Hills and Campbell (1968). The virus spreads in Australia and the UK (Garrett and O’Loughlin 1997). The virus-infected cauliflower plants exhibit symptomless infections. The virus is transmitted by an aphid vector Brevicoryne brassicae in a persistent, propagative manner (Tomlinson et al. 1972). The virus is also transmitted by mechanical sap-inoculation. The virus is not transmitted by seed. For more details of BNYV, refer to Brassica oleracea var. italica.

Cauliflower mosaic virus Taxonomic position Genus: Caulimovirus

(CaMV)

Family: Caulimoviridae

Geographical distribution CaMV infection was first reported in plants of Brassica oleracea var. botrytis from the USA by Tompkins (1937). The virus is probably distributed worldwide (Bahar 1984; Alhubaishi et al. 1987; Farzadfar et al. 2005, 2007a, b, 2008; Tuzlali and Korkmaz 2014). Symptoms and host(s) The virus-infected cauliflower plants initially exhibit symptoms on leaves which are hard to see in the field. Later in the season, mosaic patterns are easier to observe, and veinal chlorosis (vein-clearing) becomes prominent; plants are stunted and heads produced are of a small size and poor quality. Only a

Brassica oleracea var. botrytis (Cauliflower)

303

few strains of CaMV, such as NY8153, W260, D4, S-Japan, and B29, are known to infect solanaceous plant species (Piqu et al. 1995). Transmission The virus is transmitted by aphid vectors, viz., Brevicoryne brassicae and Myzus persicae, and at least 25 other species in a semi-persistent manner (Blanc et al. 2001). The virus is lost by the vector when it molts, does not multiply in the vector, is not transmitted congenitally to the progeny of the vector, but transmitted by mechanical sap-inoculation. The virus is transmitted by seed and pollen. Virion properties and genome The virions are isometric, non-enveloped, 50 nm in diameter, and rounded in profile, without a conspicuous capsomere arrangement. The capsid is constituted of 420 protein subunits organized following T = 7 icosahedral symmetry (Cheng et al. 1992). The genome is a circular double-stranded DNA molecule of 8024 bp (V00141, NC_001497) (Franck et al. 1980). The dsDNA molecule contains three gaps, two in one strand and one in the other. The gap-1 is due to the absence of one or two nucleotides with respect to the complementary beta strand. In the two gaps on the beta strand, no nucleotides are missing; instead a short sequence overlaps (19 residues for gap 2, at least 2 residues for gap 3). The genome consists of seven major open reading frames (ORF I to VII) which are all located on the () DNA strand (Guilley et al. 1982; Farzadfar et al. 2007a, 2008; Hohn 2011; Scholthof et al. 2011).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Brassica oleracea var. botrytis was reported from Yemen (Alhubaishi et al. 1987). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Radish mosaic virus Taxonomic position Genus: Comovirus

(RaMV)

Family: Secoviridae

RaMV infection in plants of Brassica oleracea var. botrytis was reported from Iran (Farzadfar et al. 2004). The virus-infected cauliflower plants exhibit mosaic, mottling, and crinkling symptoms. The virus is transmitted by beetle vectors and also by mechanical sap-inoculation, and by grafting. For more details of RaMV, refer to Raphanus sativus.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

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Brassica oleracea var. botrytis (Cauliflower)

TuMV infection in plants of Brassica oleracea var. botrytis was reported from India (Baruah and Chowfla 1989). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TuMV, refer to Brassica rapa.

Turnip yellow mosaic virus Taxonomic position Genus: Tymovirus

(TYMV)

Family: Tymoviridae

TYMV infection in plants of Brassica oleracea var. botrytis was reported from Europe and Australia (Blok et al. 1987). The virus-infected cauliflower plant symptoms begin as vein-clearing, but later and on older leaves, permanent yellow patches develop. The virus is transmitted by flea beetle vectors Phyllotreta and Psylloides spp. in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of TYMV, refer to Brassica rapa.

Turnip yellows virus Taxonomic position Genus: Polerovirus

(TuYV)

Family: Luteoviridae

TuYV infection in plants of Brassica oleracea var. botrytis was reported from China (Xiang et al. 2009, 2011). The virus-infected cauliflower plants exhibit yellowing or leaf roll symptoms. The virus is transmitted by a number of aphid species in a circulative and non-propagative manner. The virus is not mechanically sap-transmissible. For more details of TuYV, refer to Brassica rapa.

References Alhubaishi AA, Walkey DGA, Webb MJW, Bolland CJ, Cook AA (1987) A survey of horticultural plant virus diseases in the Yemen Arab Republic. FAO Plant Prot Bull 35:135–143 Bahar M (1984) Infection of cauliflower by Cauliflower mosaic virus in Isfahan. 8th Iranian congress of plant protection. Isfahan, Iran. 114 p Baruah BP, Chowfla SC (1989) Relationship of cauliflower isolate of Turnip mosaic virus with its vector, Brevicoryne brassicae L. Indian J Virol 5:56–60 Blanc S, Hebrard E, Drucker M, Froissart R (2001) Molecular basis of vector transmission: Cauliflower mosaic virus Cabb-S stran and S delta II hybrid by two species of aphid: Myzus persicae (sulzer) and Brevicoryne brassicae (L.). Res Virol 141:677–683 Blok J, Mackenzie A, Guy P, Gibbs A (1987) Nucleotide sequence comparisons of Turnip yellow mosaic virus isolates from Australia and Europe. Arch Virol 97:283–295 Cheng RH, Olson NH, Baker TS (1992) Cauliflower mosaic virus: a 420 subunit (T=7), multilayer structure. Virology 16:655–668 Farzadfar S, Pourrahim R, Golnaraghi AR, Jalali S, Ahoonmanesh A (2004) Occurrence of Radish mosaic virus on cauliflower and turnip crops in Iran. Plant Dis 88:909 Farzadfar SH, Pourrahim R, Golnaraghi AR, Ahoonmanesh A (2005) Occurrence of Cauliflower mosaic virus in different cruciferous plants in Iran. Plant Pathol 54:810 Farzadfar S, Ahoonmanesh A, Mosahebi GH, Ohshima K, Koohi-Habibi M, Pourrahim R, Golnaraghi AR (2007a) Partial biological and molecular characterization of Cauliflower mosaic virus isolates in Iran. Plant Pathol J 6:291–298 Farzadfar SH, Ahoomanesh A, Mosahebi GH, Pourrahim AR, Golnaraghi AR (2007b) Occurrence and distribution of Cauliflower mosaic virus on cruciferous plants in Iran. Plant Pathol J 6:22–29 Farzadfar SH, Mosahebi GH, Ahounmanesh A, Kouhi HD, Ohshima K, Pourrahim R, Golnaraghi AR (2008) Distribution and some biological and molecular properties of Cauliflower mosaic virus isolates from cauliflower fields in Iran. Appl Entomol Phytopathol 75(84):1–25

Brassica oleracea var. capitata (Cabbage)

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Franck A, Guilley H, Jonard G, Richards K, Hirth L (1980) Nucleotide sequence of Cauliflower mosaic virus DNA. Cell 21:285–294 Garrett RG, O’Loughlin GT (1997) Broccoli necrotic yellows virus in cauliflower and in the aphid, Brevicoryne brassicae L. Virology 76:653–663 Guilley H, Dudley RK, Jonard G, Balazs E, Richards KE (1982) Transcription of Cauliflower mosaic virus DNA: detection of promoter sequences, and characterization of transcripts. Cell 30:763–773 Hills GJ, Campbell RN (1968) Morphology of Broccoli necrotic yellows virus. J Ultrastruct Res 24:134–141 Hohn T (2011) Caulimovirus. Caulimoviridae. In: The Springer index of viruses. Springer, New York, pp 271–277. https://doi.org/10.1007/978-0-387-95919-1_41 Piqu M, Mougeot J-L, Geldreich A, Guidasci T, Mesnard J-M, Lebeurier G, Yot P (1995) Sequence of a Cauliflower mosaic virus strain infecting solanaceous plants. Gene 155:305–306 Scholthof K-BG, Adkins S, Czosnek H, Palukaitis P, Jacquot E, Hohn T, Hohn B, Saunders K, Candresse T, Ahlquist P, Hemenway C, Foster AGD (2011) Top 10 plant viruses in molecular plant pathology. Mol Plant Pathol 12:938–954 Tomlinson JA, Webb MJW, Faithfull EM (1972) Studies on Broccoli necrotic yellows virus. Ann Appl Biol 71:127–134 Tompkins CM (1937) A transmissible mosaic disease of cauliflower. J Agric Res 55:33–46 Tuzlali HT, Korkmaz S (2014) Identification and characterization of Cauliflower mosaic virus (CaMV) isolates in Canakkale province. Akdeniz Univ Ziraat Fak Derg 27:1–7 Xiang H, Shang Q, Han C, Li D, Yu J (2009) Detection of Turnip yellows virus in eight cruciferous crops in Mainland China. Phytopathology 99:S144 Xiang H-Y, Dong S-W, Shang Q-X, Zhou C-J, Wei Li D, Yu J-L, Han C-G (2011) Molecular characterization of two genotypes of a new Polerovirus infecting brassicas in China. Arch Virol 156:2251–2255

Brassica oleracea var. capitata (Cabbage) Family: Brassicaceae

Vegetable

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

BWYV causes tipburn in the heads of stored white cabbage cv. Impala (Brassica oleracea var. capitata) in the UK, Spain, and Australia (Hunter et al. 2002; Latham et al. 2003; Moreno et al. 2004). The virusinfected cabbage plants exhibit internal necrosis symptoms. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmitted by mechanical sap-inoculation. For more details of BWYV, refer to Beta vulgaris.

Cabbage cytorhabdovirus 1 (CCyV-1) Taxonomic position CCyV-1 is tentative member of the genus Cytorhabdovirus and family Rhabdoviridae Geographical distribution CCyV-1 infection in plants of Brassica oleracia var. capitata was reported from United Kingdom (Pecman et al. 2017). Transmission The virus is transmitted by the leafhopper vectors.

B

306

Brassica oleracea var. capitata (Cabbage)

Virion properties and genome The virions are bullet-shaped. The genome is a monopartite, negative sense, single-stranded RNA of 12949 nt (KY810772) (Walker et al. 2018).

Cabbage leaf curl Jamaica virus Taxonomic position Genus: Begomovirus

(CabLCJV)

Family: Geminiviridae

Geographical distribution CabLCJV infection in plants of Brassica oleracea var. capitata was reported from Jamaica (Smith 2005). Transmission The transmission of CabLCJV has not been investigated. It is likely that, in common with other begomoviruses, CabLCJV will be transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Virion properties and genome The structure of CabLCJV particles has not been investigated. In common with all geminiviruses the virions of CabLCJV will likely be geminate (twinned icosahedra). The genome of CabLCJV is bipartite consisting of two circular, single stranded DNA components. DNA-A contains 2575 nt (DQ178608) and DNA-B of 2494 nt (DQ178609). Typical of all bipartite begomoviruses native to the New World the DNA A component of CabLCJVencodes five genes, one in the virion-sense and four in the complementary-sense whereas the DNA B encodes one gene in each orientation (Briddon 2001; Smith 2005; Brown et al. 2015; Zerbini et al. 2017).

Cabbage leaf curl virus Taxonomic position Genus: Begomovirus

(CabLCV)

Family: Geminiviridae

Geographical distribution CabLCV infection in plants of Brassica oleracea var. capitata was reported from the USA (Florida and Georgia), Mexico, and Jamaica (Strandberg et al. 1991; Abouzid et al. 1992; Mandal et al. 2001; Smith 2005). Symptoms and host(s) The virus-infected cabbage plants exhibit leaf curl, interveinal chlorosis, and severe stunting symptoms. The virus has a wide host range within the Brassicaceae including cabbage, cauliflower, and Arabidopsis thaliana. Transmission The virus is transmitted by the whitefly Bemisia tabaci. It is likely that, in common with other begomoviruses, CabLCV is transmitted by B. tabaci in a circulative, non-propagative manner.

Brassica oleracea var. capitata (Cabbage)

307

The virus has been vector transmitted to cabbage/collard/cauliflower (Brassica oleracea), mustard (Brassica juncea), Chinese cabbage/turnip (Brassica rapa), radish (Raphanus raphanistrum), Rhynchosia minima, Arabidopsis thaliana, and an unidentified Desmodium spp. (Strandberg et al. 1991). Virion properties and genome The structure of the virions of CabLCV has not been investigated. In common with all geminiviruses the virions of CabLCV will likely be geminate (twinned icosahedra). CabLCV is typical of bipartite begomoviruses native to the New World. The genome consists of two circular, single-stranded DNA components. DNA-A contains 2583 nt (U65529 = NC_003866) and DNA-B of 2513 nt (U65530 = NC_003887). Typical of the majority of bipartite begomoviruses native to the New World the DNA A component of CabLCV encodes six genes, two in the virion-sense and four in the complementary-sense whereas the DNA B encodes one gene in each orientation (Abouzid et al. 1992; Hill et al. 1998; Briddon 2001; Trejo-Saavedra et al. 2009; Brown et al. 2015; Zerbini et al. 2017).

Cauliflower mosaic virus Taxonomic position Genus: Caulimovirus

(CaMV)

Family: Caulimoviridae

CaMV infects Brassica oleracea var. capitata in the UK, Iran, Spain, and Turkey (Moreno et al. 2004; Farzadfar et al. 2007; Tuzlali and Korkmaz 2014). The virus-infected cabbage plants exhibit symptoms of necrotic spots on outer leaves (Hunter et al. 2002). The virus is transmitted by aphid vectors in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of CaMV, refer to Brassica oleracea var. botrytis.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV infection in plants of Brassica oleracea var. capitata was reported from Iran (Farahani et al. 2014). The virus-infected cabbage plants exhibit symptoms of mosaic and malformations on young leaves. No definite insect is identified as a vector. The virus is mechanically sap-transmissible. The virus also spreads through mechanical transmission by field workers. For more details of TMV, refer to Nicotiana tabacum.

Tobacco necrosis virus A Taxonomic position Genus: Alphanecrovirus

(TNV-A)

Family: Tombusviridae

TNV-A infection in plants of Brassica oleracea var. capitata was reported from Lithuania (Zitikaite and Staniulis 2005). The virus-infected cabbage plants exhibit symptoms of various spotting or mottling

B

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Brassica oleracea var. capitata (Cabbage)

symptoms on leaves or fruits and necrotic symptoms on older leaves. The virus is transmitted by the fungal vector Olpidium brassicae and also by mechanical sap-inoculation. For more details of TNV-A, refer to Nicotiana tabacum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Brassica oleracea var. capitata was reported from Israel (Gera et al. 2000). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Turnip mosaic virus

(TuMV)

Synonyms Cabbage black ringspot virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

TuMV infection in plants of Brassica oleracea var. capitata occurs throughout the USA, Europe, South Africa, Asia, Japan, Australia, and Turkey (Walkey and Webb 1978; Chivasa et al. 2002; Hunter et al. 2002; Korkmaz et al. 2008; Singh et al. 2015). The virus-infected cabbage young leaves exhibits chlorotic ringspots. As the leaf ages, the ringspots are followed by yellow or brownish spots surrounded by circular or irregular necrotic rings (lesions). Generally, these occur in the vicinity of the leaf veins. Leaf blight occurs in infected sections when many lesions coalesce. TuMV also causes mosaic, black speckling, or stippling of cabbage heads at harvest or during storage. All Brassica species are susceptible to this virus. The virus is transmitted by aphid vectors Myzus persicae and Brevicoryne brassicae and other aphid species in a non-persistent manner, and also by mechanical sap-inoculation, but with no seed transmission observed. For more details of TuMV, refer to Brassica rapa.

Turnip yellows virus Taxonomic position Genus: Polerovirus

(TuYV)

Family: Luteoviridae

TuYV infection in plants of Brassica oleracea var. capitata was reported from China and South Africa (Xiang et al. 2009, 2011; New et al. 2016). The virus-infected cabbage plants exhibit yellowing or leaf roll symptoms. The virus is transmitted by a number of aphid species in a circulative and nonpropagative manner. The virus is not mechanically sap-transmissible. For more details of TuYV, refer to Brassica rapa.

Brassica oleracea var. capitata (Cabbage)

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References Abouzid AM, Hiebert E, Strandberg JO (1992) Cloning, identification and partial sequencing of a new geminivirus infecting Brassicaceae. Phytopathology 82:1070 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Chivasa S, Ekpo EJA, Hicks RGT (2002) New hosts of Turnip mosaic virus in Zimbabwe. Plant Pathol 51:386 Farahani AA, Rakhshandehroo F, Shahraeen N (2014) First report of Tobacco mosaic virus infecting cabbage in Iran. J Plant Pathol 96:S4.115 Farzadfar SH, Ahoomanesh A, Mosahebi GH, Pourrahim AR, Golnaraghi AR (2007) Occurrence and distribution of Cauliflower mosaic virus on cruciferous plants in Iran. Plant Pathol J 6:22–29 Gera A, Kritzman A, Cohen J, Raccah B, Antignus Y (2000) Tospoviruses infecting vegetable crops in Israel. EPPO Bull 30:289–292 Hill JE, Strandberg JO, Hiebert E, Lazarowitz SG (1998) Asymmetric infectivity of pseudorecombinants of Cabbage leaf curl virus and Squash leaf curl virus: implications for bipartite geminivirus evolution and movement. Virology 250:283–292 Hunter PJ, Jones JE, Walsh JA (2002) Involvement of Beet western yellows virus, Cauliflower mosaic virus, and Turnip mosaic virus in internal disorders of stored white cabbage. Phytopathology 92:816–826 Korkmaz S, Tomitaka Y, Onder S, Ohshima K (2008) Occurrence and molecular characterization of Turkish isolates of Turnip mosaic virus. Plant Pathol 57:1155–1162 Latham LJ, Smith LJ, Jones RAC (2003) Incidence of three viruses in vegetable brassica plantings and associated wild radish weeds in south-west Australia. Australas Plant Pathol 32:387–391 Mandal B, Langston DB, Pappu HR, Beard GH, Kucharek TA, Flanders JT, Whiddon JP, Smith JE, Kelley WT (2001) First report of Cabbage leaf curl virus (Family Geminiviridae) in Georgia. Plant Dis 85:561 Moreno A, de Blas C, Biurrun R, Nebreda M, Palacios I, Duque M, Fereres A (2004) The incidence and distribution of viruses infecting cultivated lettuce, cultivated brassica and associated natural vegetation in Spain. Ann Appl Biol 144:339–346 New S-A, van Heerden W, Pietersen G, Esterhuizen LL (2016) First report of a Turnip yellows virus in association with the brassica stunting disorder in South Africa. Plant Dis 100:2341 Pecman A, Kutnjak D, Gutierrez-Aguirre I, Adams I, Fox A, Boonham N, Ravnikar M (2017) Next Generation Sequencing for Detection and Discovery of Plant Viruses and Viroids: Comparison of Two Approaches. Front Microbiol 8, 1998; PMCID: PMC5645528, PMID: 29081770 Singh R, Banerjee A, Sharma SK, Bhagawati R, Baruah S, Ngachan SV (2015) First report of Turnip mosaic virus occurrence in cole crops (Brassica spp.) from Arunachal Pradesh, India. Virus Dis 26:211–213 Smith KN (2005) Molecular characterization, recombination and distribution of Cabbage leaf curl virus from Jamaica. University of the West Indies (Mona, Jamaica), M Phil thesis Strandberg JO, Hiebert E, Leibee GL, Abouzid A (1991) A new geminivirus with a broad host range in the Brassicaceae. Phytopathology 81:1244 Trejo-Saavedra DL, Vielle-Calzada JP, Rivera-Bustamante RF (2009) The infective cycle of Cabbage leaf curl virus (CaLCuV) is affected by crumpled leaf (CRL) gene in Arabidopsis thaliana. Virol J 6:169 Tuzlali HT, Korkmaz S (2014) Identification and characterization of Cauliflower mosaic virus (CaMV) isolates in Canakkale province. Akdeniz Univ Ziraat Fak Derg 27:1–7 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Rhabdoviridae. J Gen Virol 99:447–448 Walkey DGA, Webb MJW (1978) Internal necrosis in stored white cabbage caused by Turnip mosaic virus. Ann Appl Biol 89:435–441 Xiang H, Shang Q, Han C, Li D, Yu J (2009) Detection of Turnip yellows virus in eight cruciferous crops in Mainland China. Phytopathology 99:S144 Xiang H-Y, Dong S-W, Shang Q-X, Zhou C-J, Wei Li D, Yu J-L, Han C-G (2011) Molecular characterization of two genotypes of a new Polerovirus infecting brassicas in China. Arch Virol 156:2251–2255 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133 Zitikaite I, Staniulis J (2005) Isolation and characterization of Tobacco necrosis virus detected on some vegetable species. Biologia 55:35–39

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Brassica oleracea var. gemmifera (Brussels sprouts)

Brassica oleracea var. gemmifera (Brussels sprouts) Family: Brassicaceae

Turnip mosaic virus Taxonomic position Genus: Potyvirus

Vegetable

(TuMV)

Family: Potyviridae

TuMV infection in plants of Brassica oleracea var. gemmifera was reported from Turkey (Korkmaz et al. 2007, 2008). The virus-infected brussels sprout plants exhibit symptoms of concentric ringspots that occur around initial spots of infections; such spots merge together and become elongated and surrounded by necrotic rings or ring parts to form arches. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

References Korkmaz S, Onder S, Tomitaka Y, Ohshima K (2007) First report of Turnip mosaic virus on Brassicaceae crops in Turkey. Plant Pathol 56:719 Korkmaz S, Tomitaka Y, Onder S, Ohshima K (2008) Occurrence and molecular characterization of Turkish isolates of Turnip mosaic virus. Plant Pathol 57:1155–1162

Brassica oleracea var. italica (Broccoli) Family: Brassicaceae

Vegetable

Broccoli necrotic yellows cytorhabdovirus Taxonomic position Genus: Cytorhabdovirus

(BNYV)

Family: Rhabdoviridae

Geographical distribution BNYV infection in plants of Brassica oleracea var. italica was reported from England, Victoria, North America, and Australia (Lin and Campbell 1972; Tomlinson et al. 1972; Garrett and Martindale 1973). Symptoms and host(s) The virus-infected broccoli plants show mild vein-clearing and slight leaf rolling but later become symptomless. The natural host range of this virus is cauliflower, headed broccoli, and brussels sprouts.

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Transmission The virus is transmitted by an aphid vector Brevicoryne brassicae in a persistent, propagative manner (Tomlinson et al. 1972). The virus is mechanically sap-transmissible to a few plant species, Datura stramonium, which produces chlorotic to necrotic local lesions and Nicotiana glutinosa, which produces sometimes local lesions and occasionally systemic mosaics. The virus is not seed-transmitted.

Virion properties and genome The virions are enveloped and bacilliform and measure 64 nm in diameter and 297 nm long. There are distinct surface projections (spikes) dispersed evenly over all the surface. The genome is a monopartite, negative-sense, single-stranded RNA (Hills and Campbell 1968; Lin and Campbell 1972; Toriyama and Peters 1981; Dietzgen 2011; Walker et al. 2018).

Cauliflower mosaic virus Taxonomic position Genus: Caulimovirus

(CaMV)

Family: Caulimoviridae

CaMV infection in plants of Brassica oleracea var. italica was reported from Iran and Spain (Moreno et al. 2004; Farzadfar et al. 2007). The virus-infected broccoli plants exhibit mosaic spot symptoms. The virus is transmitted by aphid vectors in a semi-persistent manner, and also by mechanical sapinoculation. For more details of CaMV, refer to Brassica oleracea var. botrytis.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Brassica oleracea var. italica was reported from Turkey and Australia (Wilson et al. 2012; Sevik 2017). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Brassica oleracea var. italica was reported from India (Singh et al. 2015). The virus-infected broccoli plants exhibit mosaic, mottling, interveinal chlorosis, irregular chlorotic patches, and puckering symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

B

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Brassica oleracea var. sabauda (Kale)

References Dietzgen RG (2011) Cytorhabdovirus. Rhabdoviridae. In: The Springer index of viruses. Springer, New York, pp 1709–1713. https://doi.org/10.1007/978-0-387-95919-1_277 Farzadfar SH, Ahoomanesh A, Mosahebi GH, Pourrahim AR, Golnaraghi AR (2007) Occurrence and distribution of Cauliflower mosaic virus on cruciferous plants in Iran. Plant Pathol J 6:22–29 Garrett RG, Martindale WL (1973) The occurrence of Broccoli necrotic yellows virus in Victoria. Aust Plant Pathol Soc Newsl 2(3):17–18 Hills GJ, Campbell RN (1968) Morphology of Broccoli necrotic yellows virus. J Ultrastruct Res 24:134–141 Lin MT, Campbell RN (1972) Characterization of Broccoli necrotic yellows virus. Virology 48:30–40 Moreno A, de Blas C, Biurrun R, Nebreda M, Palacios I, Duque M, Fereres A (2004) The incidence and distribution of viruses infecting cultivated lettuce, cultivated brassica and associated natural vegetation in Spain. Ann Appl Biol 144:339–346 Sevik MA (2017) Natural occurrence of Cucumber mosaic virus infecting broccoli in Turkey. Virus Dis 28:218–219 Singh R, Banerjee A, Sharma SK, Bhagawati R, Baruah S, Ngachan SV (2015) First report of Turnip mosaic virus occurrence in cole crops (Brassica spp) from Arunachal Pradesh, India. Virus Dis 26:211–213 Tomlinson JA, Webb MJW, Faithfull EM (1972) Studies on Broccoli necrotic yellows virus. Ann Appl Biol 71:127–134 Toriyama S, Peters D (1981) Differentiation between Broccoli necrotic yellows virus and Lettuce necrotic yellows virus by their transcriptase activities. J Gen Virol 56:59–66 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Rhabdoviridae. J Gen Virol 99:447–448 Wilson CR, Lambert SJ, Dann AL, Cross P, Hay FS (2012) Occurrence of viruses within Tasmanian vegetable crops and identification of a novel Polerovirus infecting pea. Austr Plant Pathol 41:311–319

Brassica oleracea var. sabauda (Kale) Family: Brassicaceae

Vegetable

Cole latent virus

(CoLV)

Taxonomic position Genus: Carlavirus

Family: Betaflexiviridae

Geographical distribution CoLV was first reported from the state of San Paulo, Brazil, infecting cole (Brassica oleracea var. sabauda) (Kitajima et al. 1970). The virus spreads in Brazil (Belintani et al. 2002; Belintani and Gaspar 2003). Symptoms and host(s) The virus-infected plants do not exhibit any symptoms (Belintani and Gaspar 2003). Transmission The virus is transmitted by the aphid vector Myzus persicae in a non-persistent manner, and also by mechanical sap-inoculation to a number of hosts. Virion properties and genome The virions are filaments, 650 nm long, and 15 nm wide. The genome consists of a single molecule of positive-sense ssRNA of 8.3 Kb, which is polyadenylated at the 30 end and capped at the 50 end. Two

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polyadenylated subgenomic RNAs (2.6 and 1.3 Kb) are produced in CoLV-infected plants. The partial genome sequence of CoLV (30 -terminus fragment, 1040 nt) containing the coat protein, and 11.0 kDa protein is available (AY340584) (Belintani et al. 2002; Adams et al. 2004).

B References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Belintani P, Gaspar JO (2003) Coat protein and RNAs of Cole latent virus are not present within chloroplasts of Chenopodium quinoa infected cells. Fitopatol Bras 28:084–088 Belintani P, Gaspar JO, Targon MLPN, Machado MA (2002) Evidence supporting the recognition of Cole latent virus as a distinct Carlavirus. J Phytopathol 150:330–333 Kitajima EW, Camargo IJB, Costa AS (1970) Morfologia e aspectos intracelulares do vírus latente da couve. Bragantia 29:181–190

Brassica rapa (Turnip) Family: Brassicaceae

Vegetable

Ageratum enation virus Taxonomic position Genus: Begomovirus

(AEV)

Family: Geminiviridae

AEV infection in plants of Brassica rapa was reported from Pakistan (Tahir et al. 2015). The virusinfected turnip plants show foliar vein yellowing symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The virus is not mechanically saptransmissible. For more details of AEV, refer to Ageratum spp.

Beet western yellows virus

(BWYV)

Synonyms Turnip mild yellows virus Taxonomic position Genus: Polerovirus

Family: Luteoviridae

BWYV infection in plants of Brassica rapa was reported from Illinois (USA) and Israel (Marco 1984; Timmerman et al. 1985). The virus-infected turnip plants exhibit chlorosis and stunting symptoms. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. For more details of BWYV, refer to Beta vulgaris.

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Cauliflower mosaic virus Taxonomic position Genus: Caulimovirus

(CaMV)

Family: Caulimoviridae

CaMV infects Brassica rapa in the USA (Melcher 1989) and Iran (Farzadfar et al. 2005a, 2007). The virus-infected turnip plants exhibit symptoms of chlorotic spots, mottling, fine vein-clearing, necrotic spots, stunting, and rugosity. The disease symptoms depend on the age of plant and the type of CaMV isolate. The virus is transmitted by aphid vectors in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of CaMV, refer to Brassica oleracea var. botrytis.

Citrus exocortis viroid Taxonomic position Genus: Pospiviroid

(CEVd)

Family: Pospiviroidae

CEVd infection in plants of Brassica rapa was reported from Spain (Fagoaga and Duran-Vila 1996). The viroid is mechanically sap-transmissible and also through contaminated tools. For more details of CEVd, refer to Citrus spp.

Potato virus X

(PVX)

Taxonomic position Genus: Potexvirus

Family: Alphaflexiviridae

PVX was reported to infect Brassica rapa plants in India (Samad et al. 1991). The virus-infected turnip plants exhibit mosaic mottling, deformation, and distortion of leaves with overall stunting of the plant and poor roots. No insect vector is reported. The virus is mechanically sap-transmissible and has a very wide host range. The virus is transmissible by contact between plants. For more details of PVX, refer to Solanum tuberosum.

Radish mosaic virus Taxonomic position Genus: Comovirus

(RaMV)

Family: Secoviridae

RaMV infection in plants of Brassica rapa was reported from Morocco, Yugoslavia, and Iran (Stefanac and Mamula 1971; Koenig and Fisher 1981; Farzadfar et al. 2004). The virus-infected turnip plants exhibit mosaic, mottling, and crinkling symptoms. The virus is transmitted by beetle vectors and also by mechanical sap-inoculation, and by grafting. For more details of RaMV, refer to Raphanus sativus.

Brassica rapa (Turnip)

Ribgrass mosaic virus Taxonomic position Genus: Tobamovirus

315

(RMV)

Family: Virgaviridae

RMV infection in plants of Brassica rapa was reported from China and Austria (Sheng et al. 1983; Mamula and Juretic 1985). The virus-infected turnip plants exhibit mosaic, necrosis, and dwarfing symptoms. No vector is reported for this virus. The virus is mechanically sap-transmissible. The virus is transmissible by grafting and by contact between plants. For more details of RMV, refer to Plantago lanceolata.

Turnip crinkle virus

(TCV)

Taxonomic position Genus: Betacarmovirus

Family: Tombusviridae

Geographical distribution TCV infection in plants of Brassica rapa spreads in the UK, Yugoslavia, Scotland, India, and England (Broadbent and Heathcote 1958; Verma and Varma 1959; Altenbach and Howell 1981; Li and Simon 1990). Symptoms and host(s) The disease is characterized by crinkling of leaves. The infected leaves show a rugose appearance, develop yellow patches, and are brittle. The yellow patches coalesce and become necrotic with age of the plants. In the course of about a month, the affected leaves begin to die and wither away. The growth of the diseased plants is severely arrested and they present a rosette appearance (Verma and Varma 1959). The main natural hosts of this virus are turnip and other types of Brassica rapa. Transmission The virus is transmitted by flea beetle vectors Phyllotreta spp. (nine species) and Psylloides spp. (two species), in a non-persistent manner. The virus is easily transmissible by sap and contact between plants. Virion properties and genome The virions are isometric, 30 nm in diameter, and have T = 3 icosahedral symmetry. The genome consists of a single molecule of linear positive-sense ssRNA of 4054 nt (M22445 = NC_003821) and contains three open reading frames (ORFs) (Altenbach and Howell 1981; Carrington et al. 1989; Li and Simon 1990).

Turnip curly top virus Taxonomic position Genus: Turncurtovirus

(TCTV)

Family: Geminiviridae

B

316

Brassica rapa (Turnip)

Geographical distribution TCTV infection in plants of Brassica rapa was reported from Iran (Briddon et al. 2010; Razavinejad et al. 2013; Kamali et al. 2016). Symptoms and host(s) The virus-infected turnip plants exhibit leaf curling and swelling of veins on the undersides of leaves (Razavinejad and Heydarnejad 2013). The virus has been isolated from turnip (Brassica rapa var. rapa), basil (Ocimum basilicum), lettuce (Lactuca sativa), radish (Raphanus sativus), and sugar beet (Beta vulgaris) (Razavinejad et al. 2013). Transmission The virus is transmitted by the leafhopper vectors Circulifer haematoceps and Orosius albicinctus (Razavinejad and Heydarnejad 2013). The mechanism of transmission was not investigated but is likely to be circulative and non-propagative, in common with all other geminiviruses. Experimentally the virus has also been shown to infect cowpea (Vigna unguiculata) (Razavinejad et al. 2013). Virion properties and genome The structure of TCTV particles has not been investigated. In common with all geminiviruses, the virions of TCTV will likely geminate (twinned icosahedra). The genome of TCTV consists of a single, circular single-stranded DNA of 2981 nt (GU456685 = NC_014324) (Briddon et al. 2010). The characterized genomes of TCTV encode six genes - two in the virion-sense and four in the complementary-sense (Varsani et al. 2014; Kamali et al. 2017; Zerbini et al. 2017). The expression and function of the genes of TCTV has not been investigated.

Turnip leaf roll virus Taxonomic position Genus: Turncurtovirus

(TuLRV)

Family: Geminiviridae.

Geographical distribution TuLRV infection in plants of Brassica rapa was reported from Iran (Kamali et al. 2016). Symptoms and host(s) The virus-infected turnip plants exhibit leaf curling and swelling of veins on the undersides of leaves. The virus has been isolated from turnip (Brassica rapa var. rapa), lettuce (Lactuca sativa), sea beet (Beta vulgaris ssp. maritima), and sugar beet (Beta vulgaris). Transmission The virus is transmitted by the leafhopper vector Circulifer haematoceps in a circulative, non-propagative manner. Experimentally the host range has been shown to extend to radish (Raphanus sativus), rapeseed (Brassica napus), thale cress (Arabidopsis thaliana), and Nicotiana benthamiana through agroinoculation.

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Virion properties and genome The structure of TuLRV particles has not been investigated. In common with all geminiviruses, the virions of TuLRV will likely geminate (twinned icosahedra). The genome of TuLRV consists of a single, circular single-stranded DNA of 2965 nt (KT388076 = NC_029117) (Kamali et al. 2016; Zerbini et al. 2017). The characterized genomes of TuLRV isolates encode six genes - two in the virion-sense and four in the complementary-sense. The expression and function of the genes has not been investigated. However, their functions are likely the same as the positional homologs encoded by begomoviruses and curtoviruses.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

Geographical distribution TuMV infection in plants of Brassica rapa was first reported from the USA (Gardner and Kendrick 1921). The virus is probably distributed worldwide, including Africa, Asia, Europe, Oceania, North and South America, Australia, China, and Yemen (Joshi 1977; Green and Deng 1985; Alhubaishi et al. 1987; Haq et al. 1994; Farzadfar et al. 2005b). Symptoms and host(s) The virus-infected turnip plants exhibit symptoms of mosaic mottling of leaves, reduction in leaf size, and stunting. The initial symptoms to develop when Brassica seedlings are inoculated with TuMV are chlorotic spots on inoculated leaves, mottling followed by systemic vein-clearing, mosaic and/or necrosis, leaf distortion, and often stunting. Some Brassica cultivars develop progressive necrosis of leaves, petioles, and stem with some virus isolates, leading to plant death, particularly in B. napus. TuMV has a very wide host range infecting at least 318 species in 156 genera of 43 families (Edwardson and Christie 1991). Transmission The virus is transmitted by over 40 aphid species particularly by Myzus persicae, M. euphorbiae, Brevicoryne brassicae, Aphis craccivora, and A. gossypii in a non-persistent manner (Joshi 1977). Virus acquisition and inoculation take less than 1 min, there is no latent period, and aphids typically remain infective for a short period after acquiring the virus. It was shown that M. persicae retained TuMV for 3–5 h, although infectivity was lost rapidly when feeding on healthy plants. The virus is transmitted by mechanical sap-inoculation, and has a very broad host range particularly among the brassicas, and the virus is not transmitted by seed. Virion properties and genome The virions are non-enveloped, flexuous filaments, 700–750 nm long, and 11–13 nm wide. The genome is a single molecule of positive-sense ssRNA of 9835 nt (AF169561 = NC_002509) (Nicolas and Laliberte 1992). The genome has a single open reading frame that is translated into a single large polyprotein of 3863 amino acid residues, which is processed into several individual proteins (Riechmann et al. 1992; Walsh and Jenner 2002; Ohshima et al. 2007; Revers and Garcia 2015; Wylie et al. 2017).

B

318

Turnip ringspot virus

Brassica rapa (Turnip)

(TuRSV)

Taxonomic position TuRSV is a tentative member of the genus Comovirus and family Secoviridae. Geographical distribution TuRSV infection in plants of Brassica rapa was reported from Ohio (Rajakaruna et al. 2007). Symptoms and host(s) The virus-infected turnip plants exhibit chlorotic ringspots and line pattern symptoms. Transmission The virus is mechanically sap-transmissible to plants in the Brassicaceae. Virion properties and genome The virions are icosahedral particles approximately 28 nm in diameter. The genome is bipartite, linear positive-sense, single-stranded RNA. RNA1 consists of 6082 nt (FJ712026 = NC_013218; FJ516745) and RNA2 of 3985 nt (FJ712027 = NC_013219; FJ516746) (Koloniuk and Petrzik 2009; Khandekar et al. 2009; Thompson et al. 2017).

Turnip rosette virus Taxonomic position Genus: Sobemovirus

(TRoV)

Family: Solemoviridae

Geographical distribution TRoV infection in plants of Brassica rapa was recorded in Scotland and Switzerland (Broadbent and Heathcote 1958; Somera and Truve 2013). Symptoms and host(s) TRoV causes necrosis of petioles and veins, with severe dwarfing, leaf twisting, and rosetting. In Chinese cabbage, the virus induces local necrotic and occasional chlorotic spots. The natural host range of this virus is restricted to Cruciferae, Compositae, Resedaceae, and Solanaceae. TRoV was shown to infect 65 Arabidopsis ecotypes (Callaway et al. 2004). The infected plant produces systemic symptoms such as distinct vein-clearing and stunting. The ecotypes Can-0 and C24 occasionally develop local lesions. Transmission The virus is transmitted by beetle vectors and also through mechanical sap-inoculation. The virus is not seed-borne. Virion properties and genome The virions have isometric morphology measuring 28 nm in diameter (Hull 1977). The genome is polycistronic positive-sense single-stranded RNA (ssRNA) molecule of 4035 nt for (AY177608) and 4086 nt for the isolate TRoV-1 (KC778720 = NC_004553). The genome includes two overlapping ORFs, ORF2a and ORF2b (Hull 1988; Somera and Truve 2013; Somera et al. 2015).

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319

Turnip vein-clearing virus Taxonomic position Genus: Tobamovirus

(TVCV)

Family: Virgaviridae

B Geographical distribution TVCV infection in plants of Brassica rapa was first reported from Oklahoma by Lartey et al. (1993). The virus spreads in the USA (Melcher 2003). Symptoms and host(s) The virus-infected turnip plants exhibit vein-clearing symptoms. The natural host range of this virus is Plantago lanceolata, P. major, Penstemon digitalis, and Arabidopsis thaliana. Transmission There is no known vector for this virus. The virus is mechanically sap-transmissible and also through contact between plants. The virus is seed-transmitted although this occurs in the absence of infection of the embryo. Virion properties and genome The virions are non-enveloped, rigid helical rods with a helical symmetry, about 18 nm in diameter and 300–310 nm in length. The genome is a monopartite, linear, positive-sense ssRNA of 6311 nt (U03387 = NC_001873). The 30 -terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G50 pppG) (Lartey et al. 1994; Zaitlin 2011; Adams et al. 2017).

Turnip yellow mosaic virus Taxonomic position Genus: Tymovirus

(TYMV)

Family: Tymoviridae

Geographical distribution TYMV infection in plants of Brassica rapa was reported from Europe and Australia (Markham and Smith 1949; Broadbent and Heathcote 1958; Matthews 1980; Mamula and Juretic 1985). Symptoms and host(s) The virus-infected turnip plants exhibit symptoms of vein chlorosis in young leaves and then later, secondary symptom of chlorotic, pale-yellow spots that later merge into yellow zones on larger portions of the leaf surface. Finally, the green chlorophyll disappears or remains only in traces. The natural host range of this virus is almost all crops in the Cruciferae; the most important vegetables include Brassica oleracea var. capitata, B. oleracea var. botrytis, B. oleracea var. gemmifera, B. oleracea var. napobrassica, B. oleracea var. sabauda, and B. rapa. Transmission The virus is transmitted by flea beetle vectors Phyllotreta alata, P. cruciferae, P. nemorum, P. undulata, etc. and Psylloides chrysocephala and P. cuprea in a semi-persistent manner. The mustard beetles Phaedon cochleariae are also active vectors. The virus is transmissible by mechanical sap-inoculation, and grafting, but is not transmitted by pollen.

320

Brassica rapa (Turnip)

Virion properties and genome The virions are non-enveloped, isometric capsids with T = 3 icosahedral symmetry, about 30 nm in diameter. The genome consists of a single molecule of positive-sense ssRNA of 6318 nt (X07441 = NC_004063; J04373, X16378) and contains three open reading frames (ORFs), of which two of them overlap. The 30 -terminus has a tRNA-like structure (Blok et al. 1987; Morch et al. 1988; Keese et al. 1989; Dreher and Bransom 1992; Martelli et al. 2002).

Turnip yellows virus Taxonomic position Genus: Polerovirus

(TuYV)

Family: Luteoviridae

Geographical distribution TuYV was first described in Belgium in 1950 (Vanderwalle 1950). The virus infection in plants of Brassica rapa was reported from Europe, Australia, England, Germany, France, Iran, China, and Serbia (Xiang et al. 2009). Symptoms and host(s) The virus-infected turnip plants exhibit symptoms of reddening of leaf margins and interveinal yellowing and reddening of leaves, which are often unnoticed or confused with the nutritional deficiencies. The natural host range of this virus is oilseed rape, other Brassica spp. (cabbage, Brussels sprouts, kale, calabrese, cauliflower, or red cabbage), lettuce, and spinach. Transmission The virus is transmitted by a number of aphid species in a circulative and non-propagative manner. Myzus persicae is the main vector of this virus (Stevens et al. 1995). The virus is not sap or seedtransmitted. Virion properties and genome The virions are 25–30 nm in diameter and hexagonal in outline and have no envelope. TuYV capsids are composed of the major capsid protein (CP) and the minor component, the readthrough protein. The genome consists of a single molecule of positive-sense ssRNA as its genome of 5641 nt (NC_003743) (Hipper et al. 2014). The 30 terminus has neither a poly(A) tract nor a tRNA-like structure. The 50 terminus has a genome-linked protein (VPg) (of about 17 kDa). The genome contains six open reading frames (ORFs).

References Adams MJ, Adkins S, Bragard C, Gilmer D, Li D, MacFarlane SA, Wong S-M, Melcher U, Ratti C, Ryu KH, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Virgaviridae. J Gen Virol 98:1999–2000 Alhubaishi AA, Walkey DGA, Webb MJW, Bolland CJ, Cook AA (1987) A survey of horticultural plant virus diseases in the Yemen Arab Republic. FAO Plant Protect Bull 35:135–143 Altenbach SB, Howell SH (1981) Identification of a satellite RNA associated with Turnip crinkle virus. Virology 112:25–33 Blok J, Mackenzie A, Guy P, Gibbs A (1987) Nucleotide sequence comparisons of Turnip yellow mosaic virus isolates from Australia and Europe. Arch Virol 97:283–295

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Briddon RW, Heydarnejad J, Khosrowfar F, Massumi H, Martin DP, Varsani A (2010) Turnip curly top virus, a highly divergent geminivirus infecting turnip in Iran. Virus Res 152:169–175 Broadbent L, Heathcote GD (1958) Properties and host range of turnip crinkle, rosette and yellow mosaic viruses. Ann Appl Biol 46:585–592 Callaway AS, George CG, Lommel SA (2004) A Sobemovirus coat protein gene complements long-distance movement of a coat protein-null Dianthovirus. Virology 330:186–195 Carrington JC, Keaton LA, Zuidema D, Hillman BI, Morris TJ (1989) The complete genome structure of Turnip crinkle virus. Virology 170:214–218 Dreher TW, Bransom KL (1992) Genomic RNA sequence of Turnip yellow mosaic virus isolate TYMC, a cDNA-based clone with verified infectivity. Plant Mol Biol 18(2):403–406 Fagoaga C, Duran-Vila N (1996) Naturally occurring variants of Citrus exocortis viroid in vegetable crops. Plant Pathol 45:45–53 Farzadfar S, Pourrahim R, Golnaraghi AR, Jalali S, Ahoonmanesh A (2004) Occurrence of Radish mosaic virus on cauliflower and turnip crops in Iran. Plant Dis 88:909 Farzadfar SH, Pourrahim R, Golnaraghi AR, Ahoonmanesh A (2005a) Occurrence of Cauliflower mosaic virus in different cruciferous plants in Iran. Plant Pathol 54:810 Farzadfar SH, Ohshima K, Pourrahim R, Golnaraghi AR, Jalali S, Ahoonmanesh A (2005b) Occurrence of Turnip mosaic virus on ornamental crops in Iran. Plant Pathol 54:261 Farzadfar SH, Ahoomanesh A, Mosahebi GH, Pourrahim AR, Golnaraghi AR (2007) Occurrence and distribution of Cauliflower mosaic virus on cruciferous plants in Iran. Plant Pathol J 6:22–29 Gardner MW, Kendrick JB (1921) Turnip mosaic. J Agric Res 22:123–124 Green SK, Deng TC (1985) Turnip mosaic virus strains in cruciferous hosts in Taiwan. Plant Dis 69:28–31 Haq QMR, Srivastava KM, Raizada RK, Singh BP, Jain RK, Mishra A, Shukla DD (1994) Biological, serological and coat protein properties of a strain of Turnip mosaic virus causing a mosaic disease of Brassica campestris and B. juncea in India. J Phytopathol 140:55–64 Hipper C, Monsion B, Bortolamiol-Becet B, Ziegler-Graff V, Brault V (2014) Formation of virions is strictly required for Turnip yellows virus long-distance movement in plants. J Gen Virol 95:496–505 Hull R (1977) The stabilization of the particles of Turnip rosette virus and of other members of the Southern bean mosaic virus group. Virology 79:58–66 Hull R (1988) The Sobemovirus group. In: Koenig R (ed) The plant viruses. Polyhedral virions with monopartite RNA genomes, vol 3. Plenum Press, New York, pp 113–146 Joshi RD (1977) Efficiency of Aphis gossypii as vector of Turnip mosaic virus and Watermelon mosaic virus. Indian Phytopathol 30:541–544 Kamali M, Heydarnejad J, Massumi H, Kvarnheden A, Kraberger S, Varsani A (2016) Molecular diversity of turncurtoviruses in Iran. Arch Virol 161:551 Kamali M, Heydarnejad J, Pouramini N, Masumi H, Farkas K, Kraberger S, Varsani A (2017) Genome sequences of Beet curly top Iran virus, Oat dwarf virus, Turnip curly top virus, and Wheat dwarf virus identified in leafhoppers. Genome Announc 5:e01674–e01616 Keese P, Mackenzie A, Gibbs A (1989) Nucleotide sequence of the genome of an Australian isolate of Turnip yellow mosaic tymovirus. Virology 172:536–546 Khandekar S, He J, Leisner S (2009) Complete nucleotide sequence of the Toledo isolate of Turnip ringspot virus. Arch Virol 154(12):1917–1922 Koenig R, Fisher HU (1981) A Moroccan Radish mosaic virus isolate from turnip. Plant Dis 65:758–760 Koloniuk I, Petrzik K (2009) Complete genome sequence of Turnip ringspot virus. Arch Virol 154(11):1851–1853 Lartey RT, Hartson DS, Pennington RE, Sherwood JL, Melcher U (1993) Occurrence of vein-clearing tobamovirus in turnip. Plant Dis 77:21–24 Lartey RT, Lane LC, Melcher U (1994) Electron microscopic and molecular characterization of Turnip vein-clearing virus. Arch Virol 138:287–298 Li XH, Simon AE (1990) Symptom intensification on cruciferous hosts by the virulent satellite RNA of Turnip crinkle virus. Phytopathology 80:238–242 Mamula D, Juretic N (1985) A study of two isometric viruses infecting turnip in Austria. Phyton 25:241–251 Marco S (1984) Beet western yellows virus in Israel. Plant Dis 68:162–163 Markham R, Smith KS (1949) Studies on the virus of turnip yellow mosaic. Parasitology 39:330–342 Martelli GP, Sabanadzovic S, Abou-Ghanem Sabanadzovic N, Edwards MC, Dreher T (2002) The family Tymoviridae. Arch Virol 147(9):1837–1846 Matthews REF (1980) Turnip yellow mosaic virus. CMI/AAB Descriptions of Plant Viruses, No. 230 Melcher U (1989) Symptoms of Cauliflower mosaic virus infection in Arabidopsis thaliana and turnip. Bot Gaz 150:139–147 Melcher U (2003) Turnip vein-clearing virus, from pathogen to host expression profile. Mol Plant Pathol 4:133–140

B

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Brassica rapa ssp. pekinensis and chinensis (Chinese cabbage)

Morch MD, Boyer JC, Haenni AL (1988) Overlapping open reading frames revealed by complete nucleotide sequencing of Turnip yellow mosaic virus genomic RNA. Nucleic Acids Res 16:6157–6173 Nicolas O, Laliberte JF (1992) The complete nucleotide sequence of Turnip mosaic potyvirus RNA. J Gen Virol 73:2785–2793 Ohshima K, Tomitaka Y, Wood JT, Minematsu Y, Kajiyama H, Tomimura K, Gibbs AJ (2007) Patterns of recombination in Turnip mosaic virus genomic sequences indicate hot spots of recombination. J Gen Virol 88:298–315 Rajakaruna P, Khandekar S, Meulia T, Leisner SM (2007) Identification and host relations of Turnip ringspot virus, a novel Comovirus from Ohio. Plant Dis 91:1212–1220 Razavinejad S, Heydarnejad J (2013) Transmission and natural hosts of Turnip curly top virus. Iran J Plant Pathol 49:27–28 Razavinejad S, Heydarnejad J, Kamali M, Massumi H, Kraberger S, Varsani A (2013) Genetic diversity and host range studies of Turnip curly top virus. Virus Genes 46(2):345–353 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Riechmann JL, Lain S, Garcia JA (1992) Highlights and prospects of potyvirus molecular biology. J Gen Virol 73:1–16 Samad A, Naqvi QA, Garg ID, Thowene J-C (1991) Potato virus X – a new report from turnip (Brassica rapa L.) in India. J Phytopathol 133:65–75 Sheng FJ, Lu GC, Li DB (1983) An isolate of Ribgrass mosaic virus infecting Brassica rapa in the suburbia of Hang Zhou. J Zhejiang Agric Univ 9(1):39–46 Somera M, Truve E (2013) The genome organization of lucerne transient streak and turnip rosette sobemoviruses revisited. Arch Virol 158:673–678 Somera M, Sarmiento C, Truve E (2015) Overview on sobemoviruses and a proposal for the creation of the family Sobemoviridae. Viruses 7:3076–3115 Stefanac Z, Mamula D (1971) A strain of Radish mosaic virus occurring in turnip in Yugoslavia. Ann Appl Biol 69:229–234 Stevens M, Smith HG, Hallsworth PB (1995) Detection of the luteoviruses, Beet mild yellowing virus and Beet western yellows virus, in aphids caught in sugar-beet and oilseed rape crops, 1990–1993. Ann Appl Biol 127:309–320 Tahir M, Amin I, Haider MS, Mansoor S, Briddon RW (2015) Ageratum enation virus – a begomovirus of weeds with the potential to infect crops. Viruses 7:647–665 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Secoviridae. J Gen Virol 98:529–531 Timmerman EL, D’Arcy CJ, Splittstoesser WE (1985) Beet western yellows virus in Illinois vegetable crops and weeds. Plant Dis 69:933–936 Vanderwalle R (1950) La jaunisses des navets. Parasitica 6:111–112 Varsani A, Navas-Castillo J, Moriones E, Hernandez-Zepeda C, Idris A, Brown JK, Zerbini FM, Martin DP (2014) Establishment of three new genera in the family Geminiviridae: Becurtovirus, Eragrovirus and Turncurtovirus. Arch Virol 159:2193–2203 Verma GS, Varma JP (1959) Occurrence of a strain of Turnip crinkle virus in India. Indian J Microbiol 1:37–41 Walsh JA, Jenner CE (2002) Pathogen profile, Turnip mosaic virus and the quest for durable resistance. Mol Plant Pathol 3:289–300 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354 Xiang H, Shang Q, Han C, Li D, Yu J (2009) Detection of Turnip yellows virus in eight cruciferous crops in mainland China. Phytopathology 99:S144 Zaitlin M (2011) Tobamovirus. In: Tidona C, Darai G (eds) The Springer Index of Viruses. Springer, New York, NY, https://doi.org/10.1007/978-0-387-95919-1 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

Brassica rapa ssp. pekinensis and chinensis (Chinese cabbage) Family: Brassicaceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Vegetable

(ArMV)

Family: Secoviridae

Brassica rapa ssp. pekinensis and chinensis (Chinese cabbage)

323

ArMV infection in plants of Brassica rapa ssp. pekinensis was reported from New Zealand (Mossop et al. 1983). The virus-infected Chinese cabbage plants exhibit leaf mottling symptoms. This virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

B Broad bean wilt virus 2 Taxonomic position Genus: Fabavirus

(BBWV-2)

Family: Secoviridae

BBWV-2 infection in plants of Brassica rapa var. chinensis was reported from China (Cai et al. 2017). The virus-infected plants exhibit mosaic symptoms on leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BBWV-2, refer to Vicia faba.

Cauliflower mosaic virus Taxonomic position Genus: Caulimovirus

(CaMV)

Family: Caulimoviridae

CaMV infection in plants of Brassica pekinensis was reported from Iran (Farzadfar et al. 2007). The virus-infected Chinese cabbage plants exhibit chlorosis, mosaic, vein-clearing, and stunting symptoms. The virus is transmitted by aphid vectors in a semi-persistent manner, and also by mechanical sapinoculation. For more details of CaMV, refer to Brassica oleracea var. botrytis.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Brassica rapa ssp. pekinensis was reported from China (Liu et al. 2017). The virus-infected plants exhibit mosaic symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Ribgrass mosaic virus Taxonomic position Genus: Tobamovirus

(RMV)

Family: Virgaviridae

RMV infection in plants of Brassica rapa ssp. pekinensis was reported from Korea (Kim et al. 1993, 2010). The virus-infected Chinese cabbage plants exhibit symptoms of necrotic ringspots, necrotic streak on midribs, and malformation. No vector transmission is reported. The virus is mechanically

324

Brassica rapa ssp. pekinensis and chinensis (Chinese cabbage)

sap-transmissible and also transmissible through contact between plants. For more details of RMV, refer to Plantago lanceolata.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Brassica rapa ssp. pekinensis was reported from Taiwan (Green and Deng 1985). The virus-infected Chinese cabbage plants exhibit systemic mosaic and leaf deformation symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

Turnip ringspot virus

(TuRSV)

Taxonomic position TuRSV is a tentative member of the genus Comovirus and family Secoviridae. TuRSV infection in plants of Brassica rapa ssp. pekinensis was reported from Russia, the USA, and Taiwan (Koloniuk et al. 2008; Chen and Chang 2011; Chen et al. 2011). The virus-infected Chinese cabbage plants exhibit chlorotic ringspot and line pattern symptoms. The virus is mechanically saptransmissible. For more details of TuRSV, refer to Brassica rapa.

Turnip yellow mosaic virus Taxonomic position Genus: Tymovirus

(TYMV)

Family: Tymoviridae

TYMV infection in plants of Brassica rapa subspecies pekinensis was reported from Japan and Spain (Kirino et al. 2008; Alfaro-Fernandez et al. 2016). The virus-infected Chinese cabbage plants exhibit symptoms of bright yellow and dark green mosaic patterns. The virus is transmitted by flea beetle vectors Phyllotreta and Psylloides spp. in a semi-persistent manner, and also by mechanical sapinoculation. The virus is seed-transmitted at the level of up to 2.2% in Chinese cabbage (Kirino et al. 2008; Alfaro-Fernandez et al. 2016). For more details of TYMV, refer to Brassica rapa.

Turnip yellows virus Taxonomic position Genus: Polerovirus

(TuYV)

Family: Luteoviridae

TuYV infection in plants of Brassica pekinensis and B. chinensis was reported from China (Xiang et al. 2009, 2011). The virus-infected Chinese cabbage plants exhibit yellowing or leaf roll symptoms. The virus is transmitted by a number of aphid species in a circulative and non-propagative manner. The virus is not mechanically sap-transmissible. For more details of TuYV, refer to Brassica rapa.

Brassica rapa ssp. sylvestris (Wild turnip)

325

References Alfaro-Fernandez A, Serrano A, Tornos T, Cebrian MC, Cordoba-Selles MC, Jorda C, Font MI (2016) Turnip yellow mosaic virus in Chinese cabbage in Spain: commercial seed transmission and molecular characterization. Eur J Plant Pathol 146:433–442 Cai L, Zhang YC, Chen YS, Hong N, Xu WX (2017) First report of Broad bean wilt virus 2 infecting non-heading Chinese cabbage in China. Plant Dis 101:1559 Chen Y-K, Chang Y-S (2011) First report of Turnip ringspot virus in Field Mustard (Brassica chinensis) in Taiwan. Plant Dis 95:1036 Chen Y-K, Chang Y-S, Bau H-J (2011) First report of Turnip ringspot virus in field mustard (Brassica chinensis) in Taiwan. Plant Dis 95:1036 Farzadfar SH, Ahoomanesh A, Mosahebi GH, Pourrahim AR, Golnaraghi AR (2007) Occurrence and distribution of Cauliflower mosaic virus on cruciferous plants in Iran. Plant Pathol J 6:22–29 Green SK, Deng TC (1985) Turnip mosaic virus strains in cruciferous hosts in Taiwan. Plant Dis 69:28–31 Kim JS, Yoon MK, Lee KH, Choi HS (1993) Ribgrass mosaic tobamovirus occurred on Chinese cabbage in Korea. Korean J Plant Pathol 9:332 Kim J-S, Cho J-D, Choi H-S, Lee S-H, Choi G-S, Lee S-Y, Kim H-J, Yoon M-K (2010) Ribgrass mosaic tobamovirus occurred on chinese cabbage in Korea. Plant Pathol J 26:328–339 Kirino N, Inoue K, Tanina K, Yamazaki Y, Ohki ST (2008) Turnip yellow mosaic virus isolated from Chinese cabbage in Japan. J Gen Plant Pathol 74:331–334 Koloniuk I, Spak J, Petrzik K (2008) Turnip ring spot virus recognised on Chinese cabbage in Russia. Eur J Plant Pathol 122:447–450 Liu S, Cao X, Yuan X (2017) First report of Cucumber mosaic virus and its associated satellite RNA in celery cabbage in Shandong Province of China. Plant Dis 101:1829 Mossop DW, Fry PR, Young BR (1983) New plant disease records in New Zealand: Arabis mosaic virus in celery, lettuce and Chinese cabbage; Tomato spotted wilt virus in celery. NZ J Agric Res 26:257–259 Xiang H, Shang Q, Han C, Li D, Yu J (2009) Detection of Turnip yellows virus in eight cruciferous crops in Mainland China. Phytopathology 99:S144 Xiang H-Y, Dong S-W, Shang Q-X, Zhou C-J, Wei Li D, Yu J-L, Han C-G (2011) Molecular characterization of two genotypes of a new polerovirus infecting brassicas in China. Arch Virol 156:2251–2255

Brassica rapa ssp. sylvestris (Wild turnip) Family: Brassicaceae

Vegetable

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

BWYV infection in plants of Brassica rapa ssp. sylvestris was reported from Southern England (Pallett et al. 2002). The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmitted by mechanical sap-inoculation. For more details of BWYV, refer to Beta vulgaris.

Cauliflower mosaic virus Taxonomic position Genus: Caulimovirus

(CaMV)

Family: Caulimoviridae

CaMV infection in plants of Brassica rapa ssp. sylvestris was reported from Southern England (Pallett et al. 2002). The virus is transmitted by aphid vectors in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of CaMV, refer to Brassica oleracea var. botrytis.

B

326

Bromus spp. (Brome)

Turnip crinkle virus

(TCV)

Taxonomic position Genus: Betacarmovirus

Family: Tombusviridae

TCV infection in plants of Brassica rapa ssp. sylvestris was reported from Southern England (Pallett et al. 2002). The virus is transmitted by beetle vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TCV, refer to Brassica rapa.

Turnip rosette virus

(TRoV)

Taxonomic position Genus: Sobemovirus

Family: Solemoviridae

TRoV infection in plants of Brassica rapa ssp. sylvestris was reported from Southern England (Pallett et al. 2002). The virus is transmitted by beetle vectors and also by mechanical sap-inoculation. For more details of TRoV, refer to Brassica rapa.

Turnip yellow mosaic virus Taxonomic position Genus: Tymovirus

(TYMV)

Family: Tymoviridae

TYMV infection in plants of Brassica rapa ssp. sylvestris was reported from Southern England (Pallett et al. 2002). The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of TYMV, refer to Brassica rapa.

References Pallett DW, Thurston MI, Cortina-Borja M, Edwards ML, Alexander M, Mitchell E, Raybould AF, Cooper JI (2002) The incidence of viruses in wild Brassica rapa ssp. sylvestris in southern England. Annals Appl Biol 141:163–170

Bromus spp. (Brome) Family: Poaceae

Forage crop

Barley yellow dwarf virus

(BYDV)

Taxonomic position BYDV is a tentative member of the genus Luteovirus and family Luteoviridae.

Bromus spp. (Brome)

327

BYDV infection in plants of Bromus spp. was reported from western France (Henry and Dedryver 1991). The virus is transmitted by a number of aphid species in a circulative, non-propagative manner. No mechanical transmission of this virus is reported. For more details of BYDV, refer to Hordeum vulgare.

B Brome mosaic virus Taxonomic position Genus: Bromovirus

(BMV)

Family: Bromoviridae

Geographical distribution BMV was first reported in Bromus inermis from the USA by McKinney et al. (1942). The virus spreads in Europe, Australia, South Africa, and the USA (McKinney 1956; Lane 1981; Cooper 1988; Haber and Hamilton 1989; Ahlquist 1999). Symptoms and host(s) The virus-infected Bromus inermis plants exhibit light-green or with clear yellow mottling and striped spot symptoms. The natural hosts of this virus include graminaceous species, viz., Agropyron desertorum, A. repens, Avena fatua, Bromus inermis, Festuca pratensis, Hordeum murinum, Lolium multiflorum, Poa pratensis, and Setaria spp., as well as cereals, Secale cereale, Triticum aestivum, and Zea mays (Pocsai 1987; Pocsai et al. 1991). Transmission The virus is transmitted by the cereal leaf beetle vector Oulema melanopus (Gaborjanyi and Szabolcs 1987; Szabolcs and Gaborjanyi 1991). The virus is mechanically sap-transmissible to many monocotyledonous plants and a few dicotyledonous ones and usually causes a mild mosaic in most species of the Gramineae. Virion properties and genome The virions are non-enveloped about 26 nm in diameter with T = 3 icosahedral symmetry, composed of 180 capsid proteins, 12 pentamers, and 20 hexamers. The genome is segmented, tripartite linear positive-sense ssRNA composed of RNA1 (3234 nt X02380 = NC_002026), RNA2 (2865 nt X01678 = NC_002027), and RNA3 (2111 nt J02042 = NC_002028). Each genomic segment has a 30 tRNA-like structure and a 50 cap (Chiu and Sill 1963; Lane and Kaesberg 1971; Dasgupta et al. 1980; Dasgupta and Kaesberg 1982; Ahlquist et al. 1981, 1984; Damayanti et al. 1999; Choi et al. 2002; Schwartz et al. 2002; Choi and Rao 2003; Damayanti et al. 2003; Kao and Adkins 2011; Scholthof et al. 2011; Scott 2011).

Brome streak mosaic virus Synonyms Brome streak virus

(BrSMV)

328

Taxonomic position Genus: Tritimovirus

Bromus spp. (Brome)

Family: Potyviridae

Geographical distribution BrSMV infection was first reported in Bromus mollis and Hordeum murinum from the former Yugoslavia by Milicic et al. (1980, 1982). The virus spreads in the former Yugoslavia, Southern France, and Germany (Gotz and Maiss 1995). Symptoms and host(s) The virus-infected plants of Bromus mollis and Hordeum murinum exhibit chlorotic leaf streaking symptoms. Transmission The virus is transmitted by the eriophyid mite Aceria tulipae in the semi-persistent manner (Gotz and Maiss 1995). The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped and flexuous filaments, with a clear modal length of 693 nm long and 12–15 nm in diameter. The genome is a monopartite, linear, positive-sense ssRNA of 9672 nt (Z48506 = NC_003501), with a 30 -poly(A) terminus (Gotz and Maiss 1995; Lopez-Moya et al. 2009; French et al. 2011; Wylie et al. 2017).

Bromus catharticus striate mosaic virus Taxonomic position Genus: Mastrevirus

(BCSMV)

Family: Geminiviridae

Geographical distribution BCSMV infection in plants of Bromus catharticus was reported from Australia (Greber 1989; Hadfield et al. 2011). Symptoms and host(s) The virus-infected brome plants exhibit foliar striation symptoms. The virus may also occur in Paspalum dilatatum in which it also exhibits foliar striation symptoms. Transmission The virus is transmitted by the leafhopper Nesoclutha pallida (Pinner et al. 1992). In common with all other geminiviruses, BCSMV is likely transmitted in a circulative, non-propagative manner. The virus is not transmitted by mechanical inoculation. Virion properties and genome The structure of the virions of BCSMV has not been investigated. In common with all geminiviruses the virions of BCSMV will likely be geminate (twinned icosahedra) (Greber 1989; Pinner et al. 1992).

Bromus spp. (Brome)

329

The genome of BCSMV consists of a single component of circular single-stranded DNA of 2797 nt (HQ113104 = NC_014822) (Hadfield et al. 2011; Zerbini et al. 2017). In common with all mastreviruses, BCSMV is predicted to encode four gene products, two in each orientation. Based on sequence homology to previously studied mastreviruses these would code for the coat protein and movement protein in the virion-sense and in the complementary-sense for the replication-associated protein (Rep) and Rep A (Palmer and Rybicki 1998; Boulton and Davies 2011; Hadfield et al. 2011; Muhire et al. 2013).

Cocksfoot mild mosaic virus Taxonomic position Genus: Panicovirus

(CMMV)

Family: Tombusviridae

CMMV infection in plants of Bromus diandrus was reported from Europe, Canada, and New Zealand (Guy 2006). The virus-infected brome plants exhibit conspicuous mosaic symptoms. The virus is transmitted by the aphid vector Myzus persicae and also by mechanical sap-inoculation. For more details of CMMV, refer to Dactylis glomerata.

Ryegrass mottle virus Taxonomic position Genus: Sobemovirus

(RGMoV)

Family: Solemoviridae

RGMoV infection in plants of Bromus spp. was reported from Germany (Rabenstein et al. 1998). There is no known vector is reported for this virus. The virus is mechanically sap-transmissible. For more details of RGMoV, refer to Lolium spp.

References Ahlquist P (1999) Bromoviruses (Bromoviridae). In: Granoff A, Webster RG (eds) Encyclopedia of virology, vol 1, 2nd edn. Academic, San Diego, pp 198–204 Ahlquist P, Luckow V, Kaesberg P (1981) Complete nucleotide sequence of Brome mosaic virus RNA 3. J Mol Biol 153:23–38 Ahlquist P, Dasgupta R, Kaesberg P (1984) Nucleotide sequence of the Brome mosaic virus genome and its implications for viral replication. J Mol Biol 172:369–383 Boulton MI, Davies JW (2011) Mastrevirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 597–603. https://doi.org/10.1007/978-0-387-95919-1_83 Chiu RJ, Sill WH Jr (1963) Purification and properties of Bromegrass mosaic virus. Phytopathology 53:1285–1291 Choi YG, Rao AL (2003) Packaging of Brome mosaic virus RNA3 is mediated through a bipartite signal. J Virol 77:9750–9757 Choi YG, Dreher TW, Rao AL (2002) tRNA elements mediate the assembly of an icosahedral RNA virus. Proc Natl Acad Sci U S A 99:655–660 Cooper JI (1988) Brome mosaic virus (BMV). In: Smith IM et al (eds) European handbook plant diseases. Blackwell, London, p 69 Damayanti TA, Nagano H, Mise K, Furusawa I, Okuno T (1999) Brome mosaic virus defective RNAs generated during infection of barley plants. J Gen Virol 80:2511–2518

B

330

Bromus spp. (Brome)

Damayanti TA, Tsukaguchi S, Mise K, Okuno T (2003) Cis-acting elements required for efficient packaging of Brome mosaic virus RNA3 in barley protoplasts. J Virol 77:9979–9986 Dasgupta R, Kaesberg P (1982) Complete nucleotide sequences of the coat protein messenger RNAs of brome mosaic virus and cowpea chlorotic mottle virus. Nucleic Acids Res 10(2):703–713 Dasgupta R, Ahlquist P, Kaesberg P (1980) Sequence of the 30 untranslated region of brome mosaic virus coat protein messenger RNA. Virology 104(2):339–346 French RC, Stenger DC, Tatineni S (2011) Tritimovirus. Potyviridae. In: The Springer index of viruses. Springer, New York, pp 1445–1449. https://doi.org/10.1007/978-0-387-95919-1_237 Gaborjanyi R, Szabolcs J (1987) Brome mosaic virus transmission by cereal leaf beetle (Oulema melanopus, Coleoptera, Chrysomelidae). Cereal Res Commun 15:259–264 Gotz R, Maiss E (1995) The complete nucleotide-sequence and genome organization of the mite-transmitted Brome streak mosaic rymovirus in comparison with those of potyviruses. J Gen Virol 76:2035–2042 Greber R (1989) Biological characteristics of grass geminiviruses from eastern Australia. Ann Appl Biol 114:471–480 Guy PL (2006) New Zealand grasslands revisited: identification of Cocksfoot mild mosaic virus. Australas Plant Pathol 35:461–464 Haber S, Hamilton RI (1989) Brome mosaic virus isolated in Manitoba, Canada. Plant Dis 73:195–199 Hadfield J, Martin DP, Stainton D, Kraberger S, Owor BE, Shepherd DN, Lakay F, Markham PG, Greber RS, Briddon RW, Varsani A (2011) Bromus catharticus striate mosaic virus: a new mastrevirus infecting Bromus catharticus from Australia. Arch Virol 156:335–341 Henry M, Dedryver CA (1991) Occurrence of Barley yellow dwarf virus in pastures of western France. Plant Pathol 40:93–99 Kao CC, Adkins S (2011) Bromovirus. Bromoviridae. In: The Springer index of viruses. Springer, New York, pp 173–177. https://doi.org/10.1007/978-0-387-95919-1_25 Lane LC (1981) Bromoviruses. In: Kurstak E (ed) Handbook of plant virus infections and comparative diagnosis. Elesevier/North-Holland Biomedical Press, Amsterdam, pp 333–376 Lane LC, Kaesberg P (1971) Multiple genetic components in Bromegrass mosaic virus. Nat New Biol 232(28):40–43 Lopez-Moya JJ, Valli A, Garcia JA (2009) Potyviridae. In: Encyclopaedia of life sciences (ELS). Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000755.pub2 McKinney HH (1956) Interference and synergistic reactions with respects to Barley stripe mosaic virus and Brome mosaic virus. Plant Dis Reptr 40:520–523 McKinney HH, Fellows H, Johnston CO (1942) Mosaic on Bromus inermis. Phytopathology 32:331 Milicic D, Kujundzic M, Wrischer M, Plavsic B (1980) A Potyvirus isolated from Bromus mollis. Acta Bot Croat 39:27–32 Milicic D, Mamula D, Plazibat M (1982) Some properties of Brome streak mosaic virus. Acta Bot Croat 41:7–12 Muhire B, Martin DP, Brown JK, Navas-Castillo J, Moriones E, Zerbini FM, Rivera-Bustamante R, Malathi VG, Briddon RW, Varsani A (2013) A genome-wide pairwise-identity-based proposal for the classification of viruses in the genus Mastrevirus (family Geminiviridae). Arch Virol 158(6):1411–1424 Palmer KE, Rybicki EP (1998) The molecular biology of mastreviruses. Adv Virus Res 50:183–234 Pinner MS, Markham PG, Rybicki EP, Greber RS (1992) Serological relationships of geminivirus isolates from Gramineae in Australia. Plant Pathol 41:618–625 Pocsai E (1987) Effect of Brome mosaic virus infection on the plant height and weight of cereals at their early stages of growth. Cereals Res Commun 15:167–174 Pocsai E, Kobza S, Muranyi I, Szunics L (1991) Brome mosaic virus infection in different cereal breeding materials. Acta Phytopathol Entomologia Hung 26:207–212 Rabenstein F, Huth W, Lesemann DE, Ehrig F, Toriyama S (1998) First detection of Ryegrass mottle virus in Lolium breeding lines and in Bromus spec. in Germany. Book of Abstracts, 8th Conference on Virus diseases of Gramineae in Europe, 25–28 May, Goslar Scholthof K-BG, Adkins S, Czosnek H, Palukaitis P, Jacquot E, Hohn T, Hohn B, Saunders K, Candresse T, Ahlquist P, Hemenway C, Foster AGD (2011) Top 10 plant viruses in molecular plant pathology. Mol Plant Pathol 12:938–954 Schwartz M, Chen J, Janda M, Sullivan M, den Boon J, Ahlquist P (2002) A positive-strand RNA virus replication complex parallels form and function of retrovirus capsids. Mol Cell 9(3):505–514 Scott SW (2011) Bromoviridae and allies. In: Encyclopedia of life sciences (ELS). Wiley, Chichester. https://doi.org/ 10.1002/9780470015902 Szabolcs J, Gaborjanyi R (1991) Brome mosaic virus transmission by cereal leaf beetle (Oulema melanopus, Coleoptera, Chrysomelidae). Acta Phytopathol Entomol Hung 26:203–206 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

Brugmansia spp.

331

Browallia speciosa (Sapphire flower) Family: Solanaceae

Ornamental

B Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Browallia spp. was reported from the USA (Daughtrey 1996). The virusinfected sapphire flower plants exhibit symptoms of distortion, stunting, chlorosis, white or necrotic spots, stem discoloration, and/or wilt. The virus is transmitted by thrips vectors in a persistentpropagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Browallia speciosa was reported from the USA (Cho et al. 1987; Parrella et al. 2003; Hausbeck et al. 1992; Daughtrey 1996). The virus-infected sapphire flower plants exhibit symptoms of leaf chlorosis, necrotic patches, and stem discoloration. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Cho JJ, Mau RFL, Mitchell WC, Gonsalves D, Yudin LS (1987) Host list of plants susceptible to Tomato spotted wilt virus (TSWV), Research Extension Series; RES-078. University of Hawaii, Honolulu 10 p Daughtrey ML (1996) Detection and identification of tospoviruses in greenhouses. Acta Hortic 431:90–98 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among greenhouse ornamentals in Pennsylvania. Plant Dis 76:795–800 Parrella G, Gognalons P, Gebre-Selassi K, Vovlas C, Marchoux G (2003) An update of the host range of Tomato spotted wilt virus. J Plant Pathol 85:227–264

Brugmansia spp. Family: Solanaceae

Ornamental

Brugmansia mild mottle virus Taxonomic position Genus: Tobamovirus

(BrMMV)

Family: Virgaviridae

332

Brugmansia spp.

Geographical distribution BrMMV infection in Brugmansia spp. was reported from Germany and the USA (Ilmberger et al. 2007). Symptoms and host(s) The virus-infected plants exhibit mild mottling symptoms. Transmission The virus is transmissible by mechanical sap-inoculation, and also through contact between plants. Virion properties and genome The virions are rod-shaped particles with size of about 300 nm long and 18 nm wide. The genome consists of a positive-sense single-stranded RNA of 6381 nt (AM398436 = NC_010944). The 30 -terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G50 pppG) (Ilmberger et al. 2007; Zaitlin 2011; Adams et al. 2017).

Brugmansia mosaic virus Taxonomic position Genus: Potyvirus

(BruMV)

Family: Potyviridae

Geographical distribution BruMV infection in plants of Brugmansia spp. was reported from Colombia and Korea (Damsteegt et al. 2013; Zhao et al. 2013; Wylie et al 2017). Symptoms and host(s) The virus-infected Brugmansia x candida trees show mosaic, while infected B. suaveolens plants show severe mosaic, with distorted and downward-rolled leaf edges. Transmission The virus is transmitted by the aphid vector Myzus persicae in a non-persistent manner. The virus is also mechanically sap-transmissible to an extensive host range causing systemic infections in plants belonging to four different families and nine genera, with the preponderance of these belonging to the Solanaceae. Virion properties and genome The virions are non-enveloped, flexuous filaments, 720–729 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 9796 nt (JX867236 = NC_020105, JX874139) (Damsteegt et al. 2013; Zhao et al. 2013; Wylie et al. 2017).

Brugmansia suaveolens mottle virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

(BsMoV)

Brugmansia spp.

333

Geographical distribution BsMoV infection in plants of Brugmansia suaveolens was first reported from Brazil (Lucinda et al. 2008, 2010).

Symptoms and host(s) The virus-infected Brugmansia plants display leaf mottle symptoms.

Transmission The virus was transmitted by the aphid vector Myzus persicae in a non-persistent manner. The virus is also mechanically sap-transmissible.

Virion properties and genome The virions are non-enveloped, flexuous filaments, 750 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense, single-stranded RNA of 9870 nt (AB551370 = NC_014536) (Lucinda et al. 2010; Revers and Garcia 2015; Wylie et al. 2017).

Colombian datura virus Taxonomic position Genus: Potyvirus

(CDV)

Family: Potyviridae

CDV occurs in Brugmansia spp. in Colombia, the USA, Australia, Canada, India, and the UK (Lesemann et al. 1996; Adkins et al. 2005, 2008; Salamon and Palkovics 2005; Rott et al. 2009; Steele and Thomas 2009; Chellemi et al. 2011; Jordan et al. 2011; Verma et al. 2014). The virus-infected Brugmansia spp. express light and dark green mosaic symptoms, reduction in leaf size, veinbanding, and chlorotic flecking on leaves followed by mottling and plant stunting. Infection is often symptomless; however, if plants are stressed, leaf mottling and mosaic patterns will become evident. The virus is transmitted by the aphid vector Myzus persicae in a non-persistent manner. The virus is also mechanically sap-transmissible when partially purified virus is used and also through grafting. The virus spreads through the use of cuttings from the virus-infected plants for planting purpose (Salamon and Palkovics 2005). For more details of CDV, refer to Datura spp.

Potato spindle tuber viroid Taxonomic position Genus: Pospiviroid

(PSTVd)

Family: Pospiviroidae

PSTVd was reported from Brugmansia spp. in the Netherlands, Greece, the UK, Montenegro, and the Czech Republic (NPPS 2006; Verhoeven et al. 2008; Malandraki et al. 2010; Mertelik et al. 2010; Cervena et al. 2011; Matousek et al. 2014; Luigi et al. 2016). The viroid-infected Brugmansia spp. plants did not show any symptoms. The viroid is mechanically sap-transmissible. For more details of PSTVd, refer to Solanum tuberosum.

B

334

Brugmansia spp.

Tomato apical stunt viroid Taxonomic position Genus: Pospiviroid

(TASVd)

Family: Peribunyaviridae

TASVd infection in plants of Brugmansia spp. was reported from Belgium and the Netherlands (Olivier et al. 2011; Verhoeven et al. 2012). The viroid-infected Brugmansia plants do not show any symptoms. The viroid is mechanically sap-transmissible. For more details of TASVd, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Brugmansia suaveolens was reported from Korea and Serbia (Nikolic et al. 2013; Choi et al. 2014). The virus-infected Brugmansia plants exhibit symptoms of chlorotic leaf patterns and curling followed by necrosis and distortion of leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Adams MJ, Adkins S, Bragard C, Gilmer D, Li D, MacFarlane SA, Wong S-M, Melcher U, Ratti C, Ryu KH, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Virgaviridae. J Gen Virol 98:1999–2000 Adkins S, Chellemi D, Annamalai M, Baker C (2005) Colombian datura virus diagnosed in Brugmansia spp. in Florida. Phytopathology 95:52 Adkins ST, Chellemi DO, Annamalai M, Baker C (2008) Columbian datura virus re-emergence in Brugmansia spp. in the U.S. Virus Diseases of Ornamental Plants Symposium Proceedings. 21 Apr 2008 Cervena G, Necekalova J, Mikulkova H, Levkanicova Z, Mertelik J (2011) Viroids on petunia and other solanaceous crops in the Czech Republic. Acta Hortic 901:35–40 Chellemi DO, Webster CG, Baker CA, Annamalai M, Achor D, Adkins S (2011) Wide spread occurrence and low genetic diversity of Colombian datura virus in Brugmansia suggest an anthropogenic role in virus selection and spread. Plant Dis 95:755–761 Choi SK, Cho IS, Choi GS, Yoon JY (2014) First report of Tomato spotted wilt virus in Brugmansia suaveolens in Korea. Plant Dis 98:1283 Damsteegt VD, Stone AL, Smith OP, McDaniel L, Sherman DJ, Dardick C, Hammond J, Jordan R, Schneider WL (2013) A previously undescribed potyvirus isolated and characterized from arborescent Brugmansia. Arch Virol 158:1235–1244 Ilmberger N, Willingmann P, Adam G, Heinze C (2007) A subgroup 1 Tobamovirus isolated from Brugmansia spp. and its detection by RT-PCR. J Phytopathol 155:326–332 Jordan R, Guaragna MA, Putnam M (2011) Detection and molecular characterization of new and emerging potyviruses of ornamental plants. Acta Hortic 901:159–166 Lesemann D-E, Preissel H-G, Verhoeven JTJ (1996) Detection of Colombian datura potyvirus and two unidentified potyviruses in Brugmansia hybrids. Acta Hortic 432:346–353 Lucinda N, Nagata T, Inoue-Nagata AK, Salaroli RB, Kitajima EW (2008) Brugmansia suaveolens mottle virus, a novel Potyvirus causing leaf mottling of Brugmansia suaveolens in Brazil. Arch Virol 153:1971–1976 Lucinda N, Inoue-Nagata AK, Kitajima EW, Nagata T (2010) Complete genome sequence of Brugmansia suaveolens mottle virus, a Potyvirus from a ornamental shrub. Arch Virol 155:1729–1732 Luigi M, Zindovic J, Stojanovic I, Faggioli F (2016) First report of Potato spindle tuber viroid in Montenegro. J Plant Pathol 98:184

Brunfelsia spp.

335

Malandraki I, Papachristopoulou M, Vassilakos N (2010) First report of Potato spindle tuber viroid (PSTVd) in ornamental plants in Greece. New Dis Rep 21:9 Matousek J, Piernikarczyk RJJ, Dedic P, Mertelik J, Uhlirova K, Duraisamy G, Orctova L, Kloudova K, Ptacek J, Steger G (2014) Characterization of Potato spindle tuber viroid (PSTVd) incidence and new variants from ornamentals. Eur J Plant Pathol 138:93–101 Mertelik J, Kloudova K, Cervena G, Necekalova J, Mikulkova H, Levkanicova Z, Dedic P, Ptacek J (2010) First report of Potato spindle tuber viroid (PSTVd) in Brugmansia spp., Solanum jasminoides, Solanum muricatum and Petunia spp. in the Czech Republic. Plant Pathol 59:392 Nikolic D, Stankovic, Vucurovic A, Ristic D, Milojevic K, Bulajic A, Krstic B (2013) First report of Tomato spotted wilt virus on Brugmansia spp. in Serbia. Plant Dis 97:850 NPPS (Netherlands Plant Protection Service) (2006) Potato spindle tuber viroid (PSTVd) on Solanum jasminoides and Brugmansia spp. ornamental plants. Pest Report. 3 p Olivier T, Demonty E, Govers J, Belkheir K, Steyer S, Jongen C (2011) First report of a Brugmansia spp. infected by Tomato apical stunt viroid in Belgium. Plant Dis 95:495 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Rott M, Schmidt AM, Joshi V, Masters C, Godkin S, Johnson R (2009) First report of Columbian datura virus in Brugmansia in Canada. Plant Dis 93:196 Salamon P, Palkovics L (2005) Occurrence of Colombian datura virus in Brugmansia hybrids, Physalis peruviana L. and Solanum muricatum Ait. in Hungary. Acta Virol 49:117–122 Steele V, Thomas JE (2009) First report of Colombian datura virus from Australia. Aust Plant Dis Notes 4:108–109 Verhoeven JTJ, Jansen CCC, Roenhorst JW (2008) First report of pospiviroids infecting ornamentals in the Netherlands: Citrus exocortis viroid in Verbena spp., Potato spindle tuber viroid in Brugmansia suaveolens and Solanum jasminoides, and Tomato apical stunt viroid in Cestrum spp. Plant Pathol 57:399 Verhoeven JTJ, Botermans M, Meekes ETM, Roenhorst JW (2012) Tomato apical stunt viroid in the Netherlands: the most prevalent pospiviroid in ornamentals and first outbreak in tomatoes. Eur J Plant Pathol 133:803–810 Verma RK, Mishra R, Guar RK (2014) First report of Colombian datura virus in India. New Dis Rep 30:29 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354 Zaitlin M (2011) Tobamovirus. In: Tidona C, Darai G (eds) The Springer Index of Viruses. Springer, New York, NY, https://doi.org/10.1007/978-0-387-95919-1 Zhao F, Lim S, Yoo RH, Lim HS, Kwon SY, Lee SH, Moon JS (2013) Complete genome sequence of a South Korean isolate of Brugmansia mosaic virus. Arch Virol 158(9):2019–2022

Brunfelsia spp. Family: Solanaceae

Columnea latent viroid Taxonomic position Genus: Pospiviroid

Ornamental

(CLVd)

Family: Pospiviroidae

CLVd infection in plants of Brunfelsia undulata was reported from Germany (Spieker 1996). The viroid-infected Brunfelsia plants do not exhibit any symptoms. The viroid is mechanically saptransmissible. For more details of CLVd, refer to Columnea spp.

References Spieker RL (1996) A viroid from Brunfelsia undulata closely related to the Columnea latent viroid. Arch Virol 141:1823–1832

B

336

Buddleja spp. or Buddleia spp. (Butterfly bush)

Buddleja spp. or Buddleia spp. (Butterfly bush) Family: Buddlejaceae

Ornamental

Alfalfa mosaic virus

(AMV)

Taxonomic position Genus: Alfamovirus

Family: Bromoviridae

AMV infection in plants of Buddleja davidii was reported from the UK, Czechoslovakia, Germany, Italy, and France (Walter et al. 1985; Perkins and Hicks 1989; Perkins 1991). The virus-infected butterfly bush plants exhibit mosaic and leaf narrowing symptoms. The virus is transmitted by a large number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Chilli veinal mottle virus Taxonomic position Genus: Potyvirus

(ChiVMV)

Family: Potyviridae

ChiVMV infection in plants of Buddleja spp. was reported from India (Mehra et al. 2006). The virusinfected butterfly bush plants show mild chlorosis and mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ChiVMV, refer to Capsicum annuum.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Buddleja davidii was reported from the UK, Italy, the Netherlands, Eastern Germany, Yugoslavia, and France (Eric and Grbelja 1985; Perkins 1991). The virus-infected butterfly bush plants exhibit narrowing of leaf lamina along with arc and ring patterns. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Buddleja davidii was reported from the USA (Hughes and Scott 2003). The virus-infected butterfly bush plant leaves exhibited bright yellow or light green line pattern symptoms. The

Bupleurum spp.

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virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sapinoculation, and by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

References

B

Eric Z, Grbelja J (1985) Occurrence of Cucumber mosaic virus on Buddleia davidii in Yugoslavia. Acta Bot Croat 44:11–14 Hughes PL, Scott SW (2003) First report of Tomato ringspot virus in butterfly bush (Buddleia davidii). Plant Dis 87:102 Mehra A, Hallan V, Lal B, Zaidi AA (2006) Occurrence of Chilli veinal mottle virus in Himalayan butterfly bush (Buddleja crispa). Plant Pathol 55:284 Perkins CJ (1991) Virus diseases of Buddleia davidii. Plantsman 13:171–178 Perkins CJ, Hicks RGT (1989) Some properties of Alfalfa mosaic virus isolated from Buddleia davidii in the UK. Plant Pathol 38:443–446 Walter B, Kuszala J, Ravelonandro M, Pinck L (1985) Alfalfa mosaic virus isolated from Buddleia davidii compared with other strains. Plant Dis 69:266–267

Bupleurum spp. Family: Apiaceae

Medicinal

Broad bean wilt virus 2 Taxonomic position Genus: Fabavirus

(BBWV-2)

Family: Secoviridae

BBWV-2 infection in plants of Bupleurum chinense was reported from China (Sui et al. 2009). The virus-infected plants exhibit symptoms of mosaic, distortion, and stunting. The virus is transmitted by the aphid vector Acyrthosiphon solani in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BBWV-2, refer to Vicia faba.

Lettuce mosaic virus Taxonomic position Genus: Potyvirus

(LMV)

Family: Potyviridae

LMV infection in plants of Bupleurum falcatum was reported from Israel (Cohen et al. 2002). The virusinfected plants exhibit yellow mottling or mosaic and leaf distortion symptoms. The virus is transmitted by the aphid vector Myzus persicae in a non-persistent manner, and also by mechanical sap-inoculation. For more details of LMV, refer to Lactuca sativa.

References Cohen J, Rosner A, Maslenin L, Mor N, Lampel M, Zeidan M, Gera A (2002) Lettuce mosaic potyvirus is the causal agent of a new disease in Bupleurum spp. Phytoparasitica 30(1):88–95 Sui C, Wei JH, Zhan QQ, Zhang J (2009) First report of Broad bean wilt virus 2 infecting Bupleurum chinense in China. Plant Dis 93:844

338

Buxus sempervirens (Box tree)

Buxus sempervirens (Box tree) Family: Buxaceae

Tobacco streak virus Taxonomic position Genus: Ilarvirus

Trees/Shrubs

(TSV)

Family: Bromoviridae

TSV infection in plants of Buxus sempervirens was reported from the Tuscany region (Central Italy) (Bellardi et al. 2006). The virus-infected box tree plants exhibit symptoms of yellow mosaic and chlorotic stripes on the leaves and stunting of plants. The virus is transmitted by thrips vectors, is present in/on pollen, and enters into the host through the injuries caused by thrips while feeding. The virus is mechanically sap-transmissible and also by grafting. The virus is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

References Bellardi MG, Sghedoni L, Bertaccini A (2006) Tobacco streak virus infecting Buxus sempervirens. Acta Hortic 722:229–234

C

Cajanus cajan (Red gram/Pigeon pea) Family: Fabaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Grain legume

(AMV)

Family: Bromoviridae

AMV infection in plants of Cajanus cajan was reported from New Zealand (Fletcher 1989). The virusinfected pigeon pea plants exhibit interveinal mottling and chlorotic ringspot symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Mungbean yellow mosaic virus Taxonomic position Genus: Begomovirus

(MYMV)

Family: Geminiviridae

MYMV infection in plants of Cajanus cajan was reported from Sri Lanka, India, Myanmar, the Philippines, Jamaica, Nepal, and Puerto Rico (Nene et al. 1971; Reddy et al. 1990, 1998; Mandal et al. 1998). The virus-infected pigeon pea plants show conspicuous bright yellow mosaic and mottling of leaves and bear fewer pods when infected at early stage of plant growth. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of MYMV, refer to Vigna radiata.

Pigeon pea sterility mosaic emaravirus 1 Taxonomic position Genus: Emaravirus

Family: Fimoviridae

© Springer Nature India Private Limited 2019 K. S. Sastry et al., Encyclopedia of Plant Viruses and Viroids, https://doi.org/10.1007/978-81-322-3912-3

(PPSMV1)

340

Cajanus cajan (Red gram/Pigeon pea)

Geographical distribution PPSMV1 was first reported on Cajanus cajan in 1931 from India (Mitra 1931; Alam 1933). The virus spreads in India, Nepal, Bangladesh, Myanmar, Sri Lanka, and Thailand (Capoor 1952; Reddy et al. 1990; Nene 1995; Singh et al. 1999; Kulkarni 2002; Dharmaraj et al. 2004; Jones et al. 2004; Kumar et al. 2005; Patil and Kumar 2015; Surender Kumar et al. 2016). Symptoms and host(s) The virus-infected pigeon pea plants show pale green mosaic symptoms and excessive vegetative growth and fail to produce flowers (sterility) and pods. The leaves of infected plants are small with light and dark green mosaic. Mosaic symptoms initially appear as vein-clearing on young leaves. When infection occurs 45 days after emergence, only some parts of the plant show symptoms, while the remaining pods appear normal. Very late infections does not show clear symptoms, but when ratooned, the new growth shows clear mosaic symptoms and sterility (Latha 2003). The natural hosts of this virus are pigeon pea and its wild relatives (Cajanus albicans, C. cajanifolius, C. lineatus, C. platycarpus, C. scarabaeoides, and C. sericeus) (Kulkarni et al. 2003). Transmission This virus is transmitted by a mite vector Aceria cajani in a semi-persistent manner (Seth 1962; Oldfield et al. 1981). A. cajani acquired PPSMVafter a minimum acquisition access period (AAP) of 15 min and inoculated virus after a minimum inoculation access period (IAP) of 90 min. No latent period was observed. Starvation of A. cajani prior to, or following, PPSMVacquisition reduced the minimum AAP and IAP periods to 10 min and 60 min, respectively, and mites retained virus for up to 13 h. None of the mites that developed from eggs taken from PPSMV-infected leaves transmitted the virus, indicating that it is not transmitted transovarially. The virus is mechanically sap-transmissible to Nicotiana benthamiana, but not to pigeon pea (Kulkarni 2002; Kulkarni et al. 2002; Jones et al. 2004). The virus is transmitted through grafting and is not transmitted through pollen or seed (Kumar et al. 2002, 2003). PPSMVoccurs as several geographically distinct pathotypes, which differ in their severity on pigeon pea genotypes (Reddy et al. 1993; Kumar et al. 2004). Virion properties and genome The virions are quasi-spherical, double membrane-bound particles (DMBs) of 80 to 100 nm (Patil and Kumar 2015). The genome consists of five segments of negative-sense single-stranded RNA that are predicted to encode proteins. The largest segment, RNA-1, is 7022 nt (HF568801 = NC_029575); RNA-2, with a sequence length of 2223 nt (HF568802 = NC_029556); RNA-3, with a sequence length of 1442 nt (HF568803 = NC_029574); RNA-4, with a sequence length of 1563 nt (HF568804 = NC_029557); and RNA-5, with a sequence length of 1689 nt (HF945448 = NC_029569) (Kumar et al. 2003; Mielke-Ehret and Muhlbach 2012; Elbeaino et al. 2014; Patil and Kumar 2015; Surender Kumar et al. 2016; Elbeaino et al. 2018).

Pigeon pea sterility mosaic emaravirus 2 Taxonomic position Genus: Emaravirus

(PPSMV2)

Family: Fimoviridae

Geographical distribution PPSMV2 infection in plants of Cajanus cajan was reported from Italy (Elbeaino et al. 2015).

Cajanus cajan (Red gram/Pigeon pea)

341

Symptoms and host(s) The virus-infected pigeon pea plants exhibit chlorotic mottling and mild mosaic symptoms. Transmission The virus is transmitted by a mite vector, Aceria cajani. Virion properties and genome The virions are quasi-spherical, double membrane-bound particles (DMBs) of 80 to 100 nm. The genome consists of six segments of negative-sense ssRNAs. RNA-1 consists of 7009 nt (HF912243 = NC_030660), RNA-2 of 2229 nt (HF912244 = NC_030662), RNA-3 of 1335 nt (HF912245 = NC_030661), RNA-4 of 1491 nt (HF912246 = NC_030663), RNA-5 of 1833 nt (HG939489 = NC_030658), and RNA-6 of 1194 nt (HG939490 = NC_030659) (Mielke-Ehret and Muhlbach 2012; Elbeaino et al. 2015; Elbeaino et al. 2018).

Pothos latent virus Taxonomic position Genus: Aureusvirus

(PoLV)

Family: Tombusviridae

PoLV infection in plants of Cajanus cajan was reported from India (Kumar et al. 2001). The virusinfected pigeon pea plants generally do not exhibit any external symptoms. The virus is transmitted experimentally through soil to herbaceous test plants but not to pigeon pea. The virus is mechanically sap-transmissible to herbaceous hosts such as N. benthamiana, N. clevelandii, and N. hispens. For more details of PoLV, refer to Scindapsus spp.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Cajanus cajan was reported from India and Australia (Vemana et al. 2014; Sharman et al. 2015). The virus-infected pigeon pea plants exhibit symptoms of necrotic spots on young leaves followed by wilting of leaves, petiole, and branch/axillary shoot proliferation with small leaves having mosaic symptoms and stem necrosis. The virus is transmitted by the thrips vectors, the virus present in/on pollen enters into the host through the injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Cajanus cajan was reported from India, Nepal, Bangladesh, Sri Lanka, Thailand, and Myanmar (Raj et al. 2005). The virus-infected pigeon pea plants exhibit

C

342

Cajanus cajan (Red gram/Pigeon pea)

symptoms of yellow mosaic, mottling, shortening of leaves, and stunting. The virus is transmitted by Bemisia tabaci in a circulative non-propagative manner. Mechanical inoculations failed to transmit the disease. For more details of ToLCNDV, refer to Solanum lycopersicum.

References Alam M (1933) Arhar sterility. Proc Twentieth Ann Meet Indian Sci Congr Poona Sect Agric 43:15–16 Capoor SP (1952) Observations on the sterility disease of pigeonpea in Bombay. Indian J Agric Sci 22:271–274 Dharmaraj PS, Narayana YD, Kumar PL, Waliyar F, Jones AT (2004) Pigeonpea sterility mosaic disease: an emerging problem in northern Karnataka. Int Chickpea Pigeonpea Newsl 11:47–49 Elbeaino T, Digiaro M, Uppala M, Sudini H (2014) Deep sequencing of Pigeonpea sterility mosaic virus discloses five RNA segments related to emaraviruses. Virus Res 188:27–31 Elbeaino T, Digiaro M, Uppala M, Sudini H (2015) Deep-sequencing of dsRNAs recovered from mosaic-diseased pigeonpea (Cajanus cajan L.) revealed the presence of a novel Emaravirus: Pigeonpea sterility mosaic virus 2 (PPSMV2). Arch Virol 160:2019–2029 Elbeaino T, Digiaro M, Mielke-Ehret N, Muelbach H-P, Martelli GP, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Fimoviridae. J Gen Virol 99:1478–1479 Fletcher JD (1989) Additional hosts of Alfalfa mosaic virus, Cucumber mosaic virus and Tobacco mosaic virus in New Zealand. N Z J Crop Hortic Sci 17:361–362 Jones AT, Kumar PL, Saxena KB, Kulkarni NK, Muniyappa V, Waliyar F (2004) Sterility mosaic disease – the “green plague” of pigeonpea: advances in understanding the etiology, transmission and control of a major virus disease. Plant Dis 88:436–445 Kulkarni N (2002) Studies on pigeonpea sterility mosaic disease; transmission, virus-vector relationships and identification of resistant sources. PhD thesis submitted to UAS, Bangalore, India, pp 137 Kulkarni NK, Kumar PL, Muniyappa V, Jones AT, Reddy DVR (2002) Transmission of Pigeonpea sterility mosaic virus by the eriophyid mite, Aceria cajani (Acari: Arthropoda). Plant Dis 86:1297–1302 Kulkarni NK, Kumar PL, Muniyappa V, Jones AT, Reddy DVR (2003) Studies on host range of Pigeonpea sterility mosaic virus. J Mycol and Plant Pathol 33:141–145 Kumar PL, Jones AT, Sreenivasulu P, Fenton B, Reddy DVR (2001) Characterization of a virus from pigeonpea with affinities to species in the genus Aureusvirus, family Tombusviridae. Plant Dis 85:208–215 Kumar PL, Jones AT, Reddy DVR (2002) Mechanical transmission of Pigeonpea sterility mosaic virus. J Mycol Plant Pathol 32:88–89 Kumar PL, Jones AT, Reddy DVR (2003) A novel mite transmitted virus with a divided RNA genome closely associated with pigeonpea sterility mosaic disease. Phytopathology 93:71–81 Kumar PL, Jones AT, Waliyar F (2004) Biology, etiology and management of pigeonpea sterility mosaic disease. Ann Rev Plant Pathol 3:1–24 Kumar PL, Latha TKS, Chauhan VB, Aparna V, Waliyar F (2005) Studies on Pigeonpea sterility mosaic virus isolates in India. In: Abstracts second Asian conference on plant pathology, National University of Singapore. pp 85–86 Kumar S, Subba Rao BL, Hallan V (2016) Virome analysis of sterility mosaic disease infected pigeonpea by small RNA deep sequencing. In: IVS international conference on “Global Perspectives in Virus Disease Management”, VIROCON 2016, December 8–10th, Bangalore, India, p 99. (OPPV18) Latha TKS (2003) Studies on pigeonpea sterility mosaic disease: characterization of Coimbatore isolate and identification of resistant sources. PhD (Ag.) thesis submitted to Tamil Nadu Agriculture University, Coimbatore, Tamil Nadu, India Mandal B, Varma A, Malathi VG (1998) Some biological properties of pigeonpea isolates of Mungbean yellow mosaic geminivirus. Indian Phytopathol 51:121–129 Mielke-Ehret N, Muhlbach HP (2012) Emaravirus: a novel genus of multipartite, negative strand RNA plant viruses. Viruses 4:1515–1536 Mitra M (1931) Report of imperial mycologist. Sci Rept Imp Inst Agric Res Pusa 1929–30:45–71 Nene YL (1995) Sterility mosaic of pigeonpea: the challenge continues. Indian J Mycol Plant Pathol 25:1–11 Nene YL, Naresh JS, Nair NG (1971) Additional host of Mungbean yellow mosaic virus. Indian Phytopathol 24:415–417 Oldfield GN, Reddy MV, Nene YL, Reed W (1981) Preliminary studies of the eriophyid vector of sterility mosaic. Int Chickpea Pigeonpea Newsl 1:25–27 Patil BL, Kumar PL (2015) Pigeonpea sterility mosaic virus: a legume-infecting Emaravirus from South Asia. Mol Plant Pathol 16(8):775–786 Raj SK, Khan MS, Singh R (2005) Natural occurrence of a Begomovirus on pigeonpea in India. Plant Pathol 54:809 Reddy MV, Sharma SB, Nene YL (1990) Pigeon pea: disease management. In: Nene YL, Hall SD, Sheila VK (eds) The Pigeonpea. CAB International, Wallingford, pp 303–347

Caladenia spp. (C. arenicola; C. latifolia) (Spider orchid)

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Reddy MV, Raju TN, Nene YL, Ghanekar AM, Amin KS, Arjunan G, Astaputre JV, Sinha BK, Reddy SV, Gupta RP, Gangadharan K (1993) Variability in sterility mosaic pathogen in pigeonpea in India. Indian Phytopathol 46:206–212 Reddy MV, Raju TN, Lenne JM (1998) Diseases of pigeonpea. In: Allen DJ, Lenne JM (eds) The pathology of food and pasture legumes. CAB International, Wallingford, pp 517–558 Seth ML (1962) Transmission of pigeonpea sterility by an eriophyid mite. Indian Phytopathol 15:225–227 Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct Tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207 Singh AK, Rathi YPS, Agrawal KC (1999) Sterility mosaic of pigeonpea: a challenge of the 20th century. Indian J Virol 15:85–92 Vemana K, Seshadri Goud TE, Reddy DL, Venkateswarlu NC, Naik KSS, Sampath Kumar D, Padmalatha Y, Desai S (2014) First report of Tobacco streak virus infecting pigeon pea (Cajanus Cajan) in India. Plant Dis 98:287

Cajanus scarabaeoides Family: Fabaceae

Medicinal plant

Pigeonpea sterility mosaic emaravirus 1 Taxonomic position Genus: Emaravirus

(PPSMV1)

Family: Fimoviridae

PPSMV infection in plants of Cajanus scarabaeoides was reported from India (Reddy et al. 1993). The virus-infected plants show pale green mosaic symptoms. This virus is transmitted by a mite vector, Aceria cajani, in a semi-persistent manner. For more details of PPSMV, refer to Cajanus cajan.

References Reddy MV, Sheila VK, Nene YL (1993) Cajanus scarabaeoides an alternate host of pigeon pea sterility mosaic pathogen and its vector Aceria cajani. Int Pigeonpea Newsl 18:24–26

Caladenia spp. (C. arenicola; C. latifolia) (Spider orchid) Family: Orchidaceae

Ornamental

Caladenia virus A

(CalVA)

Taxonomic position Genus: Poacevirus

Family: Potyviridae

Geographical distribution CalVA infection in plants of Caladenia spp. was reported from Australia (Wylie et al. 2012).

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Caladium spp.

Symptoms and host(s) The virus-infected spider orchid plants exhibit mild to severe streaking symptoms. Transmission A mite is the possible vector of this virus. Virion properties and genome The virions are non-enveloped, flexuous filaments, and 680–750 nm long and 12–15 nm in diameter. The genome is a monopartite, linear, positive-sense ssRNA of 9847 nt (JX156425 = NC_018572) with a 30 -poly(A) terminus (Wylie et al. 2012, 2017).

Donkey orchid symptomless virus Taxonomic position Genus: Platypuvirus

(DOSV)

Family: Alphaflexiviridae

DOSV infection in plants of Caladenia spp. was reported from Western Australia (Wylie et al. 2013). The virus-infected spider orchid plants do not exhibit any symptoms. The virus is mechanically saptransmissible. For more details of DOSV, refer to Diuris spp.

References Wylie SJ, Tan AJY, Li H, Dixon KW, Jones MGK (2012) Caladenia virus A, an unusual new member of the family Potyviridae from terrestrial orchids in Western Australia. Arch Virol 157:2447–2452 Wylie SJ, Li H, Jones MGK (2013) Donkey orchid symptomless virus: a viral ‘platypus’ from Australian terrestrial orchids. PLoS One 8:e79587 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Caladium spp. Family: Araceae

Caladium virus X

Ornamental

(CalVX)

Taxonomic position CalVX is a tentative member of the genus Potexvirus and family Alphaflexiviridae Geographical distribution CalVX infection in plants of Caladium spp. was reported from Brazil (Rivas et al. 2005). Symptoms and host(s) The virus-infected plants exhibit systemic chlorotic spots, green rings, and mosaic symptoms on leaves.

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Transmission The virus is mechanically sap-transmissible to 13 species that belongs to the families Amaranthaceae, Araceae, Chenopodiaceae, Plantaginaceae, Portulacaceae, and Solanaceae. The virus is also transmissible through contact between plants and through implements and workers’ hands. The virus is not transmitted by aphid vectors or through seeds. Virion properties and genome The virions are slightly flexuous rods measuring 420–479 nm length and 13 nm in diameter. The genome is linear unipartite positive-sense single-stranded RNA (Rivas et al. 2005). A partial genome sequence of 738 nt is available (AY727533).

Dasheen mosaic virus Taxonomic position Genus: Potyvirus

(DsMV)

Family: Potyviridae

DsMV infection in plants of Caladium bicolor was reported from Brazil, Western Samoa, and the USA (Hartman and Zettler 1974; Van Velsen 1979; Nelson 2008). The virus-infected plants exhibit foliar mosaic symptoms which often appear as a localized “feathering” of white tissue along the veins on several cultivars. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of DsMV, refer to Colocasia esculenta.

Konjac mosaic virus Taxonomic position Genus: Potyvirus

(KoMV)

Family: Potyviridae

KoMV infection in plants of Caladium spp. was reported from Japan, Taiwan, and India (Padmavathi et al. 2011). The virus-infected plants exhibit symptoms of mosaic, chlorotic feathery mottle, chlorotic spots or ringspots, and leaf malformation. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of KoMV, refer to Amorphophallus paeoniifolius.

References Hartman RD, Zettler FW (1974) Effects of Dasheen mosaic virus on yields of caladium, dieffenbachia and philodendron. Phytopathology 64:768 Nelson SC (2008) Dasheen mosaic of edible and ornamental aroids. Plant Disease CTAHR. Available at: http://www. ctahr.hawaii.edu/oc/freepubs/pdf/PD-44.pdf Padmavathi M, Srinivas KP, Subba Reddy CV, Ramesh B, Navodayam K, Krishnaprasadji J, Babu Ratan P, Sreenivasulu P (2011) Konjac mosaic virus naturally infecting three aroid plant species in Andhra Pradesh, India. J Phytopathol 159:133–135 Rivas EB, Duarte LML, Alexandre MAV, Galleti SR, Harakava R, Fernandes FMC (2005) Caladium virus X, a new potexvirus from Caladium bicolor (Araceae). J Plant Pathol 87:109–114 Van Velsen RJ (1979) Final report on the UNDP/FAO survey of plant viruses in Western Samoa. UNDP/FAO-SPEC Survey of agricultural pests and diseases of the South Pacific. 19 pp

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Calanthe spp.

Calanthe spp. Family: Orchidaceae

Ornamental

Bean yellow mosaic virus Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Calanthe spp. was reported from Japan and the USA (Inouye and Inouye 1972; Hammond and Lawson 1988; Matsumoto et al. 1999). The virus-infected plants exhibit symptoms of chlorotic spots on the leaves. The virus is transmitted by an aphid vector, Myzus persicae, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Calanthe mild mosaic virus Taxonomic position Genus: Potyvirus

(CalMMV)

Family: Potyviridae

Geographical distribution CalMMV infection in Calanthe orchids was reported from Japan (Gara et al. 1998). Symptoms and host(s) The virus-infected plants show a mild mosaic and flower break symptoms. Transmission The virus is transmitted by an aphid vector, Myzus persicae in a non-persistent manner. The virus is also mechanically sap-transmitted to Calanthe spp. and Phalaenopsis spp. Virion properties and genome The virions are non-enveloped and flexuous filaments 764 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense, single-stranded RNA. A partial genome sequence of 1306 nt is available (AB011404) (Gara et al. 1998; Revers and Garcia 2015; Wylie et al. 2017).

Clover yellow vein virus Taxonomic position Genus: Potyvirus

(ClYVV)

Family: Potyviridae

ClYVV was identified in Japan from Calanthe discolor and C. sieboldii showing mosaic and/or veinal necrosis on the foliage and floral necrosis (Inouye et al. 1988). The virus is transmitted by an aphid vector Myzus persicae in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ClYVV, refer to Trifolium spp.

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347

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV was reported from wild populations of endemic Calanthe on an isolated island in Japan (Inouye et al. 1982; Kawakami et al. 2008). The virus-infected plants exhibit mild mosaic and etching symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Odontoglossum ringspot virus Taxonomic position Genus: Tobamovirus

(ORSV)

Family: Virgaviridae

ORSV infection in plants of Calanthe spp. was reported from Japan (Matsumoto et al. 1995). The virusinfected plants exhibit chlorotic mottle symptoms on the leaves. The virus is transmitted by means not involving a vector. The virus is transmissible by mechanical sap-inoculation, and also through contact between plants. For more details of ORSV, refer to Odontoglossum grande.

Orchid fleck dichorhavirus Taxonomic position Genus: Dichorhavirus

(OFV)

Family: Rhabdoviridae

OFV infection in plants of Calanthe spp. was reported from Japan (Inouye et al. 1996). The virusinfected plants exhibit symptoms of light green and/or yellowish fleck mosaic on the leaves. The virus is transmitted by mite vectors in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of OFV, refer to Cymbidium spp.

References Gara IW, Kondo H, Maeda T, Tamada T, Inouye N (1998) Calanthe mild mosaic virus, a new Potyvirus from Calanthe orchid in Japan. J Phytopathol 146:357–363 Hammond J, Lawson RH (1988) A strain of Bean yellow mosaic virus is aphid-transmitted from orchid. Acta Hortic 234:365–370 Inouye N, Inouye T (1972) On the Bean yellow mosaic virus isolated from Calanthe orchid. Ann Phytopathol Soc Jpn 38:211 Inouye N, Maeda T, Mitsuhata K (1982) Cucumber mosaic virus isolated from Calanthe discolor. Nogaku Kenkyu 60:1–11 Inouye N, Maeda T, Mitsuhata K (1988) A strain of Clover yellow vein virus isolated from Calanthe sp. Acta Hortic 234:61–68 Inouye N, Matsumoto J, Maeda T, Mitsuhata K, Kondo H, Tahara M (1996) Orchid fleck virus, the causal agent of a yellowish fleck mosaic disease of Calanthe. Bull Res Inst Bioresour Okayama Univ 4:119–135 Kawakami K, Fuji S-I, Miyoshi K (2008) Endangered wild populations of endemic Calanthe orchids on an isolated Japanese island tested for viruses. Aust J Bot 55:831–836 Matsumoto J, Maeda T, Inouye N (1995) Odontoglossum ring spot virus isolated from Calanthe sp. in Japan. Bull Res Inst Bioresour Okayama Univ 3:163–174 Matsumoto JI, Maeda T, Inouye N (1999) Some properties of Bean yellow mosaic virus isolated from Calanthe sp. orchidaceae in Japan. Bull Res Inst Bioresour Okayama Univ 6(1):43–51 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

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Calceolaria spp.

Calceolaria spp. Family: Calceolariaceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Calceolaria spp. was reported from the USA (Daughtrey 1996). The virusinfected plants exhibit symptoms of leaf chlorosis, brown necrotic spots with lighter centers, irregular brown spotting, leaf necrosis, stem discoloration, and stunting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Calceolaria spp. was reported from the USA (Cho et al. 1987; Hausbeck et al. 1992; Daughtrey 1996; Parrella et al. 2003). The virus-infected plants exhibit symptoms of leaf necrosis along the main veins. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Cho JJ, Mau RFL, Mitchell WC, Gonsalves D, Yudin LS (1987) Host list of plants susceptible to Tomato spotted wilt virus (TSWV), Research Extension Series; RES-078. University of Hawaii, Honolulu. 10 p Daughtrey ML (1996) Detection and identification of tospoviruses in greenhouses. Acta Hortic 431:90–98 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among greenhouse ornamentals in Pennsylvania. Plant Dis 76:795–800 Parrella G, Gognalons P, Gebre-Selassi K, Vovlas C, Marchoux G (2003) An update of the host range of Tomato spotted wilt virus. J Plant Pathol 85:227–264

Calendula spp. Family: Asteraceae

Ageratum enation virus Taxonomic position Genus: Begomovirus

Ornamental

(AEV)

Family: Geminiviridae

Calendula spp.

349

AEV infection in plants of Calendula officinalis was reported from India (Jaidi et al. 2015). The virusinfected plants exhibit yellow vein net symptoms on the leaves and leaf deformation. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmitted by mechanical sap-inoculation. For more details of AEV, refer to Ageratum spp.

Bidens mottle virus Taxonomic position Genus: Potyvirus

C

(BiMoV)

Family: Potyviridae

BiMoV was identified from a high percentage of field-grown pot marigold (Calendula officinalis) at one location in Taiwan (Huang and Jan 2011); the infected plants show chlorotic spots, generalized chlorosis, and stunting. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BiMoV, refer to Bidens spp.

Cucumber mosaic virus

(CMV)

Synonyms Calendula yellow net virus Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV infection in plants of Calendula officinalis was reported from India, Serbia, and Italy (Lisa and Dellavalle 1979; Naqvi and Samad 1985; Milosevic et al. 2015). The virus-infected plants exhibit yellow netting or chlorotic mottle and leaf deformation symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV was reported from Iran in Calendula spp. (Ghotbi et al. 2005; Ghotbi 2013). The virus-infected plants were stunted, yielding a small number of twisted and deformed flowers. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Pepino mosaic virus Taxonomic position Genus: Potexvirus

(PepMV)

Family: Alphaflexiviridae

350

Calendula spp.

PepMV infection in plants of Calendula arvensis was reported from Cyprus (Papayiannis et al. 2012). The virus is mechanically sap-transmissible. For more details of PepMV, refer to Solanum muricatum.

Tobacco curly shoot virus

(TbCSV)

Synonyms Calendula yellow vein virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

TbCSV infection in plants of Calendula officinalis was reported from India (AY887174) (Khan et al. 2005, 2007). The virus-infected plants exhibit symptoms of vein yellowing, shortening of leaves and petioles, and stunting of plants. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of TbCSV, refer to Nicotiana tabacum.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV was reported in plants of Calendula officinalis as early as 1925 in the USA (Elmer 1925). The virus-infected plants exhibit mosaic symptoms. There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Calendula spp. was reported from Iran (Ghotbi 2006). The virus-infected plants exhibit mosaic, leaf deformation, and dwarfing symptoms. The virus is transmitted by the thrips vectors. The virus present in/on pollen, and enters into the host through injuries caused by the thrips while feeding. The virus is also mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato mosaic virus Taxonomic position Genus: Tobamovirus

(ToMV)

Family: Virgaviridae

ToMV was reported from Calendula officinalis in Bulgaria (Hristova et al. 1994). The virus-infected plants exhibit indistinct mosaic symptoms. No vector is involved in the spread of this virus. This virus is

Calendula spp.

351

transmissible by mechanical sap-inoculation, transmissible by grafting, and transmissible by contact between plants. For more details of ToMV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

C Family: Tospoviridae

TSWV was reported in plants of Calendula spp. in Iran and Bulgaria (Cho et al. 1987; Moini and Izadpanah 2000; Ghotbi et al. 2005; Parrella et al. 2003; Dikova 2011). The virus-infected plants exhibit symptoms of flower and leaf deformation and necrotic lesions on the leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Calendula arvensis was reported from Cyprus (Papayiannis et al. 2011). The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. The virus is transmitted by grafting but not transmitted by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Calendula officinalis was reported from Italy (Lisa and Dellavalle 1979). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of TuMV, refer to Brassica rapa.

References Cho JJ, Mau RFL, Mitchell WC, Gonsalves D, Yudin LS (1987) Host list of plants susceptible to Tomato spotted wilt virus (TSWV), Research Extension Series; RES-078. University of Hawaii, Honolulu. 10 p Dikova B (2011) Tomato spotted wilt virus on some medicinal and essential oil-bearing plants in Bulgaria. Bulg J Agric Sci 17:306–313 Elmer OH (1925) Transmissibility and pathological effects of the mosaic disease. Iowa State Agric Exp Station Res Bull 82:55 Ghotbi T (2006) First report on incidence of Tobacco streak virus (TSV) on ornamental plants in Iran. Iran J Plant Pathol 42:159–160 Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iran J Plant Pathol 49:41

352

Calibrachoa spp.

Ghotbi T, Sharaeen N, Winter S (2005) Occurrence of tospoviruses in ornamental and weed species in Markazi and Tehran provinces in Iran. Plant Dis 89:425–429 Hristova D, Barkerdzhieva N, Svrakov K (1994) Tomato mosaic virus isolated from Calendula officinalis. Plant Protection Institute, 2230 Kostinbrod, Bulgaria, Int Conf Pl Virol. Apriltsi, Troyon, Bulgaria 32:153 Huang CH, Jan FJ (2011) First report of Bidens mottle virus infecting calendula in Taiwan. Plant Dis 95:362 Jaidi M, Kumar S, Srivastava A, Raj SK (2015) First report of Ageratum enation virus and ageratum leaf curl betasatellite infecting Calendula officinalis in India. New Dis Rep 36:6 Khan AA, Naqvi QA, Khan MS, Singh R, Raj SK (2005) First report of a Begomovirus infecting calendula in India. Plant Pathology 54:569 Khan AA, Khan MS, Raj SK, Naqvi QA (2007) Molecular identification of a Begomovirus causing yellow vein disease on Calendula officinalis in India. EPPO Bull 37:420–426 Lisa V, Dellavalle G (1979) Isolation of two viruses from Calendula officinalis. Inf Fitopathol 29:11–12 Milosevic D, Ignjatov M, Nikolic Z, Gvozdanovic-Varga J, Petrovic G, Stankovic I, Krstic B (2015) First report of Cucumber mosaic virus causing chlorotic mottle on pot marigold (Calendula officinalis) in Serbia. Plant Dis 99(5):736 Moini AA, Izadpanah K (2000) New hosts for Tomato spotted wilt virus in Tehran. Iran J Plant Pathol 36:104–105 Naqvi QA, Samad A (1985) Purification and properties of Calendula yellow net virus. Indian J Virol 1:143–146 Papayiannis L, Katis NI, Idris AM, Brown JK (2011) Identification of weed hosts of Tomato yellow leaf curl virus in Cyprus. Plant Dis 95:120–125 Papayiannis LC, Kokkinos CD, Alfafo-Fernandez A (2012) Detection, characterization and host range studies of Pepino mosaic virus in Cyprus. Eur J Plant Pathol 132:1–7 Parrella G, Gognalons P, Gebre-Selassi K, Vovlas C, Marchoux G (2003) An update of the host range of Tomato spotted wilt virus. J Plant Pathol 85:227–264

Calibrachoa spp. Family: Solanaceae

Ornamental

Calibrachoa mottle virus Taxonomic position Genus: Alphacarmovirus

(CMoV)

Family: Tombusviridae

Geographical distribution CMoV infection in plants of Calibrachoa spp. was reported from the USA (Liu et al. 2003) and also known to be present in Europe and Asia. Symptoms and host(s) The virus-infected plants show streaking in the flowers, interveinal chlorosis, and mottling or blotching on the leaves. Some cultivars will not show symptoms unless they are stressed. Transmission The virus is mechanically sap-transmitted to Chenopodium amaranticolor, C. capitatum, C. quinoa, Nicotiana benthamiana, and N. clevelandii. The virus is not transmitted by the aphid vector, Myzus persicae, or any of several species of whiteflies (Liu et al. 2003). Virion properties and genome The virions are isometric 30 nm in diameter and have T = 3 icosahedral symmetry. The genome consists of a single molecule of linear positive-sense ssRNA of 3919 nt (GQ244431 = NC_021926) (GulatiSakhuja and Liu 2010).

Callistephus spp.

353

Tobacco mild green mosaic virus Taxonomic position Genus: Tobamovirus

(TMGMV)

Family: Virgaviridae

TMGMV was reported in plants of Calibrachoa spp. from Israel and the USA (Gera et al. 2007; Zeidan et al. 2008; Sabanadzovic et al. 2009). The virus-infected Calibrachoa spp. exhibited chlorotic yellow spots, rings, vein-banding, and mosaic on the foliage, and the plants exhibited reduced growth. In some cultivars this virus caused general chlorosis and blistering of foliage. No insect vector is reported for this virus. The virus is transmissible by mechanical sap-inoculation, and has a wide host range. The virus is transmissible by grafting and also by contact between plants. For more details of TMGMV, refer to Nicotiana tabacum.

References Gera A, Beckelman H, Lipsker Z, Maslenin L, Spiegel S, Ziedan M (2007) Identification of Tobacco mild green mosaic virus in Torenia, Petunia and Calibrachoa. Olam Haperach, Sept–Oct, pp 58–60 Gulati-Sakhuja A, Liu HY (2010) Complete nucleotide sequence and genome organization of Calibrachoa mottle virus (CbMV) – a new species in the genus Carmovirus of the family Tombusviridae. Virus Res 147:216–223 Liu HY, Sears JL, Morrison RH (2003) Isolation and characterization of a carmo-like virus from Calibrachoa plants. Plant Dis 87:167–171 Sabanadzovic S, Henn A, Abou Ghanem-Sabanadzovic N, Lawrence A (2009) First report on Tobacco mild green mosaic virus in Calibrachoa Plants (Calibrachoa  hybrida) in Mississippi. Plant Dis 93:1354 Zeidan M, Beckelman E, Holdengreber P et al (2008) Detection of Tobacco mild green mosaic virus in ornamental crops. In: Proceedings of the 12th international symposium on virus diseases of ornamental plants, Haarlem, the Netherlands, 20–24 April 2008, p 52

Callistephus spp. Family: Asteraceae

Ornamental

Callistephus chinensis chlorosis virus

(CCCV)

Taxonomic position CCCV is a tentative member of the Family Rhabdoviridae. Geographical distribution CCCV infection in plants of Callistephus chinensis was reported from the South and Central American region and Brazil (Francki et al. 1985). Symptoms and host(s) The virus-infected Callistephus chinensis plants show leaf chlorosis symptoms. Virion properties and genome The virions are rhabdo- or bullet-shaped, with a clear modal length of 200–300 nm and 60–80 nm wide. Virions contain a single negative-sense RNA of >12 kb.

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Callistephus spp.

Callistephus mottle virus Taxonomic position Genus: Potyvirus

(CalMoV)

Family: Potyviridae

Geographical distribution CalMoV infection in plants of Callistephus chinensis was reported from South Korea (Seo et al. 2016). Symptoms and host(s) The virus-infected Callistephus chinensis plants show severe foliar mottling symptoms. No other natural hosts are known. Transmission The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long and 11–13 nm wide. The genome contains single molecule of linear, positive-sense single-stranded RNA consisted of 9,859 nt (excluding the poly(A) tail) (KX013584 = NC_030794) and contains the typical open reading frame of potyviruses, encoding a putative large polyprotein of 3,154 amino acids (Seo et al. 2016; Wylie et al. 2017).

Chrysanthemum stem necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(CSNV)

Family: Tospoviridae

CSNV was detected in a plant of Callistephus spp. in Brazil, showing symptoms of mosaic, with leaf browning and stem necrosis (Alexandre et al. 1999). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of CSNV, refer to Chrysanthemum spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection has been reported in Chinese aster (Callistephus chinensis) in Korea (Oh et al. 2008). The virus-infected plants exhibit severe mosaic symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Callistephus spp.

355

Groundnut ringspot orthotospovirus Taxonomic position Genus: Orthotospovirus

(GRSV)

Family: Tospoviridae

GRSV was detected in a plant of Callistephus spp. in Brazil, showing symptoms of mosaic with necrotic spots and lines on the leaves (Alexandre et al. 1999). The virus is transmitted by thrips vectors in a persistent-propagative manner, and the virus is also mechanically sap-transmissible. For more details of GRSV, refer to Arachis hypogaea.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Callistephus chinensis was reported from the USA (Daughtrey 1996). The virus-infected plants exhibit symptoms of ringspots, stem cankers, and necrotic or chlorotic line patterns. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Callistephus chinensis was reported from the USA (Daughtrey 1996; Bakardjieva et al. 1998). The virus-infected plants show chlorotic or necrotic ringspots and leaf necrosis symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Alexandre MAV, Duarte LML, Rivas EB, Chagas CM (1999) Mixed infection of Tospovirus species in ornamental crops in Sao Paolo State, Brazil. Summa Phytopathol 25:353–356 Bakardjieva N, Denkova S, Hristova D (1998) Tomato spotted wilt virus on ornamental species in Bulgaria. Biotechnol Biotechnol Eq 12:49–52 Daughtrey ML (1996) Detection and identification of tospoviruses in greenhouses. Acta Hortic 431:90–98 Francki RIB, Milne RG, Hatta T (1985) Atlas of plant viruses, vol II. CRC Press, Boca Raton Oh SM, Kim SR, Hong JS, Ryu KH, Lee GP, Choi JK (2008) Characterization of an isolate of Cucumber mosaic virus isolated from Chinese aster (Callistephus chinensis). Res Plant Dis 14:229–232 Seo E-Y, Lim S, Hammond J, Moon JS, Lim H-S (2016) Complete genome sequence of a novel Potyvirus, Callistephus mottle virus identified in Callistephus chinensis. Arch Virol 161:3281–3283 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

C

356

Calopogonium mucunoides (Calopo)

Calopogonium mucunoides (Calopo) Family: Fabaceae

Forage crop

Calopogonium yellow vein virus Taxonomic position Genus: Tymovirus

(CalYVV)

Family: Tymoviridae

Geographical distribution CalYVV infection in plants of Calopogonium mucunoides was reported from Malaysia (Gibbs et al. 1997). Symptoms and host(s) The virus-infected calopo plants exhibit yellow vein symptoms. Transmission The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sapinoculation. Virion properties and genome The virions are non-enveloped, isometric capsid with T = 3 icosahedral symmetry, about 30 nm in diameter. The genome consists of a single molecule of positive-sense ssRNA. The 30 terminus has a tRNA-like structure. A partial genome sequence of 696 nt is available (U91413) (Gibbs et al. 1997; Martelli et al. 2002).

Cowpea severe mosaic virus Taxonomic position Genus: Comovirus

(CPSMV)

Family: Secoviridae

CPSMV infection in plants of Calopogonium mucunoides was reported from central Brazil (Lin et al. 1982). The virus-infected calopo plants exhibit symptoms of severe mosaic and blistering in the leaves. The virus is transmitted by beetle vectors and also by mechanical sap-inoculation. For more details of CPSMV, refer to Vigna unguiculata.

References Gibbs AJ, Mackenzie AM, Abdul-Samad N (1997) A tymovirus from Calopogonium mucunoides in Malaysia is not Clitoria yellow vein tymovirus. Arch Virol 142(8):1697–1702 Lin MT, Anjos JRN, Rios GP (1982) Cowpea severe mosaic virus in five legumes in central Brazil. Plant Dis 66:67–70 Martelli GP, Sabanadzovic S, Abou-Ghanem Sabanadzovic N, Edwards MC, Dreher T (2002) The family Tymoviridae. Arch Virol 147(9):1837–1846

Calycanthus floridus (Sweet shrub)

357

Calotropis spp. (Calotropis gigantea; C. procera) (Crown flower, Rubber bush) Family: Apocynaceae

Medicinal

C Groundnut bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(GBNV)

Family: Tospoviridae

GBNV infection in plants of Calotropis gigantea was reported from Andhra Pradesh (India) (Bhaskara Reddy et al. 2011). The virus-infected crown flower plants exhibit symptoms of mosaic, chlorotic rings, and necrosis. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of GBNV, refer to Arachis hypogaea.

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Calotropis procera was reported from Pakistan (Zaidi et al. 2017). The virus-affected rubber bush plants exhibit leaf yellowing and yellow mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible through grafting and not through the seed. For more details of ToLCNDV, refer to Solanum lycopersicum.

References Bhaskara Reddy BV, Sivaprasad Y, Sai Gopal DVR (2011) First report of Groundnut bud necrosis virus on Calotropis gigantea. J Plant Pathol S4.84 Zaidi SS, Shakir S, Malik HJ, Farooq M, Amin I, Mansoor S (2017) First report of Tomato leaf curl New Delhi virus on Calotropis procera, a weed as potential reservoir begomovirus host in Pakistan. Plant Dis 101:1071

Calycanthus floridus (Sweet shrub) Family: Calycanthaceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(INSV)

358

Camassia spp. (Indian hyacinth)

INSV infection in plants of Calycanthus floridus was reported from Georgia, USA (Ruter and Gitaitis 1993). The virus-infected sweet shrub plants exhibit symptomless infections. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is mechanically sap-transmissible to a large number of hosts. For more details of INSV, refer to Impatiens spp.

References Ruter JM, Gitaitis RD (1993) Impatiens necrotic spot virus in woody landscape plants in Georgia. Plant Dis 77:318

Camassia spp. (Indian hyacinth) Family: Asparagaceae

Ornamental

Arabis mosaic virus

(ArMV)

Taxonomic position Genus: Nepovirus

Family: Secoviridae

ArMV infection in plants of Camassia cusickii and C. quamash was reported from Lithuania (Samuitiene et al. 2008). The virus-infected Indian hyacinth plants exhibit symptoms of light green streaks and pinpoint necrotic lesions on leaves. The virus is transmitted by nematode vectors, Xiphinema diversicaudatum and X. coxi, in a non-persistent manner, and also by mechanical sapinoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

References Samuitiene M, Navalinskiene M, Jackeviciene E (2008) Arabis mosaic virus on ornamental plants. Biologia 54:264–268

Camelina sativa (Camelina) Family: Brassicaceae

Oil seed

Turnip yellow mosaic virus Taxonomic position Genus: Tymovirus

(TYMV)

Family: Tymoviridae

TYMV infection in plants of Camelina sativa was reported by Hein (1984). The virus-infected camelina plants exhibit symptoms of leaf spots and stunted growth. The virus is transmitted by beetle vectors in a semi-persistent manner, and also through mechanical sap-inoculation. The virus is

Campanula spp. (Bellflower)

359

seed-transmitted in Camelina sativa plants up to 20% (Hein 1984). For more details of TYMV, refer to Brassica rapa.

References Hein A (1984) Transmission of Turnip yellow mosaic virus by seed of Camelina sativa (gold of pleasure). Z Pfl Krankh Pfl Schutz 91:549–551

Campanula spp. (Bellflower) Family: Campanulaceae

Apple mosaic virus Taxonomic position Genus: Ilarvirus

Ornamental

(Acronym: ApMV)

Family: Bromoviridae

ApMV infection in plants of Campanula spp. was reported from Turkey (Arli Sokmen et al. 2005, 2008). The virus-infected bellflower plants exhibit stunting and vein-clearing symptoms. No insect vector is known for this virus. The virus is mechanically sap-transmissible and also by grafting. For more details of ApMV, refer to Malus domestica.

Bellflower vein chlorosis virus Taxonomic position Genus: Waikavirus

(BVCV)

Family: Secoviridae

Geographical distribution BVCV infection in plants of Campanula takesimana was reported from Korea (Seo et al. 2015). Symptoms and host(s) The virus-infected bellflower plants exhibit vein chlorosis symptoms. Transmission The virus is not mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is a monopartite linear, positive-sense, single-stranded RNA of 11,649 nt (KT238881 = NC_027915) (Sanfacon et al. 2009; Hibino 2011; Sanfacon 2015; Seo et al. 2015; Thompson et al. 2017).

C

360

Campanula spp. (Bellflower)

Bellflower veinal mottle virus Taxonomic position Genus: Bevemovirus

(BVMoV)

Family: Potyviridae

Geographical distribution BVMoV infection in plants of Campanula takesimana was reported from South Korea (Seo et al. 2017). Symptoms and host(s) The virus-infected bellflower plants exhibit veinal mottle symptoms on the leaves. Transmission The virus is probably not aphid transmitted because the HC-Pro lacks the conserved potyvirus aphid transmission motifs R/KITC and PTK. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long and 11–13 nm wide. The genome contains single molecule of linear, positive-sense single-stranded RNA of 8259 nt (KY491536 = NC_039002). The genome encodes a single large ORF of 7416 nt, predicted to be processed into nine mature proteins (Seo et al. 2017; Wylie et al. 2017).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV has been reported to infecting plants of Campanula grandiflora in Japan (Komuro and Asuyama 1955) and C. rapunculoides in the Czech Republic (Polak 1964). The virus-infected bellflower plants exhibit symptoms of flower streaking and distortion. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Tomato aspermy virus Taxonomic position Genus: Cucumovirus

(TAV)

Family: Bromoviridae

TAV infection in plants of Campanula rapunculoides was reported from the Netherlands (Noordam et al. 1965). The virus-infected bellflower plants exhibit symptoms of concentric rings and line patterns in leaves and some flower break. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TAV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

Campsis radicans (Trumpet vine)

361

TSWV-infected Campanula spp. plants were reported from China, the Czech Republic, and Brazil (Hausbeck et al. 1992; Mokra et al. 2002; Gioria et al. 2010; Wan et al. 2017). The virus-infected bellflower plants exhibit symptoms of mosaic, leaf necrosis, and flower ringspot. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Arli Sokmen M, Kutluk Yilmaz ND, Mennan H, Sevik MA (2005) Natural weed hosts of Apple mosaic virus in hazelnut orchards in Turkey. J Plant Pathol 87:239–242 Arli Sokmen M, Kutluk Yilmaz ND, Mennan H, Sevik MA (2008) RT-PCR detection of Apple mosaic virus infection in some weed hosts found in hazelnut orchards in Turkey. Acta Hortic 781:155–160 Gioria R, Brunelli KR, Kobori RF, Kobori MMRG, Rezende JAM, Kitajima EW (2010) First report of Tomato spotted wilt virus (TSWV) infecting Campanula medium in Brazil. Summa Phytopathol 36(2):176–177 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among greenhouse ornamentals in Pennsylvania. Plant Dis 76:795–800 Hibino H (2011) Waikavirus. Sequiviridae. In: The Springer index of Viruses. Springer, New York, pp 1775–1779. https:// doi.org/10.1007/978-0-387-95919-1_289 Komuro Y, Asuyama H (1955) Studies on Cucumber mosaic virus: II. The isolation of the virus from various plants showing mosaic in the vicinity of Tokyo, Japan. Jpn J Phytopathol 20:77–82 Mokra V, Gotzova B, Mertelik J, Polak J (2002) Collection of ornamental plant viruses. Acta Hortic 568:193–199 Noordam D, Bijl M, Overbeek SC, Quiniones SS (1965) Virusen uit Campanula rapunculoides en Stellaria media en hut relatie tot konkommermosaiek virus en tomaat-‘aspermi’-virus. Neth J Plant Pathol 71:61 Polak Z (1964) Campanula rapunculoides L. a natural source of Cucumber mosaic virus. Preslia 36:306 Sanfacon H (2015) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. In: eLS. Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000764.pub3. http://www.els.net Sanfacon H, Wellink J, Le Gall O, Karasev A, van der Vlugt R, Wetzel T (2009) Secoviridae: a proposed family of plant viruses within the order Picornavirales that combines the families Sequiviridae and Comoviridae, the unassigned genera Cheravirus and Sadwavirus, and the proposed genus Torradovirus. Arch Virol 154(5):899–907 Seo JK, Kwak HR, Lee YJ, Kim J, Kim MK, Kim CS, Choi HS (2015) Complete genome sequence of Bellflower vein chlorosis virus, a novel putative member of the genus Waikavirus. Arch Virol 160(12):3139–3142 Seo JK, Kwak HR, Kim MK, Kim JS, Choi HS (2017) The complete genome sequence of a novel virus, Bellflower veinal mottle virus, suggests the existence of a new genus within the family Potyviridae. Arch Virol 162(8):2457–2461 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J Gen Virol 98:529–531 Wan YR, Yin YQ, Wu QJ (2017) First report of Tomato spotted wilt virus infecting balloon flower in China. J Plant Pathol 99:799–818 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Campsis radicans (Trumpet vine) Family: Bignoniaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Ornamental

(AMV)

Family: Bromoviridae

C

362

Canavalia spp. (Sword bean, Jack-bean)

AMV infection in plants of Campsis radicans was reported from Iran (Pourrahim and Farzadfar 2016). The virus-infected trumpet vine plants exhibit mottling and mosaic symptoms. The virus is transmitted by a large number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

References Pourrahim R, Farzadfar S (2016) Biological and molecular characterization of Alfalfa mosaic virus infecting trumpet creeper (Campsis radicans) in Iran. J Phytopathol 164:276–280

Canavalia spp. (Sword bean, Jack-bean) Family: Fabaceae

Grain legume

Cowpea aphid-borne mosaic virus Taxonomic position Genus: Potyvirus

(CABMV)

Family: Potyviridae

CABMV infection in plants of Canavalia rosea was reported from Brazil (Kitajima et al. 2008). The virus-infected sword bean plants exhibit mosaic symptoms. The virus is transmitted by a number of aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CABMV, refer to Vigna unguiculata.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV infection in plants of Canavalia ensiformis was reported from Venezuela (Marys et al. 2004). The virus-infected sword bean plants exhibit symptoms of severe stunting, leaf mosaic, mottling, distortion, and general yellowing. There is no known vector for this virus. The virus is mechanically saptransmissible and also through contact between plants. For more details of TMV, refer to Nicotiana tabacum.

References Kitajima EW, de Alc^antara BK, Madureira PM, Alfenas-Zerbini P, Rezende JAM, Zerbini FM (2008) A mosaic of beach bean (Canavalia rosea) caused by an isolate of Cowpea aphid-borne mosaic virus (CABMV) in Brazil. Arch Virol 153(4):743–747 Marys E, Ortega E, Carballo O (2004) Natural infection of Canavalia ensiformis with Tobacco mosaic virus in Venezuela. Plant Dis 88:681

Canna spp. (Canna edulis, C. indica, C. glauca, C. generalis)

363

Canna spp. (Canna edulis, C. indica, C. glauca, C. generalis) Family: Cannaceae

Ornamental

Banana streak virus Taxonomic position Genus: Badnavirus

C

(BSV)

Family: Caulimoviridae

BSV infection in plants of Canna edulis was reported from Colombia (Reichel et al. 1997). The virusinfected canna plants exhibit mild mosaic symptoms. The virus is transmitted by mealybug vectors in a semi-persistent manner. The virus is not transmissible by mechanical sap-inoculation. For more details of BSV, refer to Musa spp.

Bean yellow mosaic virus

(BYMV)

Synonyms Canna mosaic virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

BYMV was reported from the USA (Chauhan et al. 2015) causing mosaic in Canna glauca and C. generalis and is considered the same as Canna mosaic infecting C. indica in the Philippines, India, and Russia (Brierley and Smith 1948; Castillo et al. 1956; Kumari et al. 2015; Zakubanskiy et al. 2017; Mitrofanova et al. 2018). This virus is probably present wherever cannas are grown. The virus-infected plants show severe yellow mosaic symptoms, and red coloration is quite narrow along the veins and mostly depleted from the margins. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Canna yellow mottle associated virus Taxonomic position Genus: Badnavirus

(CaYMAV)

Family: Caulimoviridae

Geographical distribution CaYMAV infection in plants Canna indica was reported from Oklahoma, USA (Wijayasekara et al. 2018). Symptoms and host(s) The virus-infected canna plants exhibit yellow mottle symptoms.

364

Canna spp. (Canna edulis, C. indica, C. glauca, C. generalis)

Transmission The virus is transmitted by mealybug vectors in a semipersistent manner. The virus is transmitted by mechanical inoculation (with difficulty). The virus is transmitted by grafting and not by contact between plants or by pollen. Virion properties and genome The virions are bacilliform in shape with a diameter of 30 nm and modal particle length of 130 nm. The genome is a circular dsDNA of 6966 bp (KX066020 = NC_030462) and encodes three open reading frames (ORFs) on the positive DNA strand (Olszewski and Lockhart 2011; Bhat et al. 2016; Wijayasekara et al. 2018).

Canna yellow mottle virus Taxonomic position Genus: Badnavirus

(CaYMV)

Family: Caulimoviridae

Geographical distribution CaYMV infection in plants of Canna spp. was reported from Russia, Austria, India, Japan, Italy, the Netherlands, the UK, the USA, and Kenya (Lockhart 1988; Momol et al. 2004; Borroto-Fernandez et al. 2008; Marino et al. 2008; Pappu et al. 2008; Windham 2008; Kumari et al. 2014; Agneroh et al. 2015; Chauhan et al. 2015; Zakubanskiy et al. 2017). Symptoms and host(s) The virus-infected canna plants show symptoms such as mottling, general yellowing, and veinal chlorosis. Transmission No vector has yet been identified. The virus is not mechanically sap-transmissible. The virus spreads through the use of infected planting materials like suckers and rhizomes. Virion properties and genome The virions are bacilliform, non-enveloped, and 120–130 nm in length and 28 nm wide. The genome consists of a single molecule of non-covalently closed circular dsDNA of c.7.1–7.4 kbp; three genomic sequences are available (KU168312 = NC_038380, 7120 bp; MF074075, 7385 bp; KY971493, 7348 bp) suggesting the possibility of two distinct badnaviruses associated with the disease in this crop (Yamashita et al. 1985; Olszewski and Lockhart 2011; Bhat et al. 2016; Wijayasekara et al. 2018).

Canna yellow streak virus Taxonomic position Genus: Potyvirus

(CaYSV)

Family: Potyviridae

Geographical distribution CaYSV infection in plants of Canna spp. was reported from the UK, Belgium, France, the Netherlands, the USA, Brazil, Russia, and Israel (Monger et al. 2007; Chauhan et al. 2015; Alexandre et al. 2017; Zakubanskiy et al. 2017; Mitrofanova et al. 2018).

Canna spp. (Canna edulis, C. indica, C. glauca, C. generalis)

365

Symptoms and host(s) The virus-infected canna plants exhibit yellow mosaic, streaking, and necrosis along the veins. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments 760 nm long and 12 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9502 nt (GQ421689 = NC_013261) (Monger et al. 2010; Revers and Garcia 2015; Wylie et al. 2017).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Canna spp. was reported from Russia, Ohio (USA), and Iran (Fisher et al. 1997; Jeon et al. 2006; Samuitiene and Navalinskiene 2008; Saidi and Safaeizadeh 2012; Zakubanskiy et al. 2017; Mitrofanova et al. 2018). The virus-infected canna plants exhibit symptoms of mosaic, chlorotic and distorted foliage, and severe stunting. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts and has been reported to be seed-transmitted in Canna up to 23% (Brito et al. 2012). For more details of CMV, refer to Cucumis sativus.

Sugarcane mosaic virus Taxonomic position Genus: Potyvirus

(SCMV)

Family: Potyviridae

SCMV infection in plants of Canna spp. was reported from China (Tang et al. 2016, 2018). The virusinfected canna plants exhibit mosaic and chlorosis symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SCMV, refer to Saccharum officinarum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Canna indica was reported from Iran (Moini and Izadpanah 2000). The virus-infected canna plants exhibit marginal necrosis of the leaves and flower deformation and wilting symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

C

366

Canna spp. (Canna edulis, C. indica, C. glauca, C. generalis)

References Agneroh TA, Bratsch SA, Lockhart BE (2015) First report of Canna yellow mottle virus in Kenya. Plant Health Prog. https://doi.org/10.1096/PHP-BR-14-0037 Alexandre MAV, Duarte LML, Chaves ALR, Ramos AF, Harakava R, Kitajima EW (2017) Canna paniculata as natural host of Canna yellow streak virus in Brazil. Aust Plant Dis Notes 12:38 Bhat AI, Hohn T, Selvarajan R (2016) Badnaviruses: the current global scenario. Viruses 8(6):177 Borroto-Fernandez EG, Maghuli F, Fellner A, Laimer M (2008) Determination of virus infections in an Austrian collection of Canna indica. J Plant Dis Protect 115:102–103 Brierley P, Smith FF (1948) Canna mosaic in the United States. Phytopathology 38:230–234 Brito M, Garrido MJ, Mejias A, Marys E (2012) Seed transmission of Cucumber mosaic virus in canna (Canna indica). Fitopatología Venez 25:24–26 Castillo BS, Yarwood CE, Gold AH (1956) Canna mosaic virus. Plant Dis Reptr 40:169–172 Chauhan RP, Hamon HF, Rajakaruna P, Webb MA, Payton M, Verchot J (2015) Reliable detection for Bean yellow mosaic virus, Canna yellow streak virus, and Canna yellow mottle virus in canna varieties with red foliage. Plant Dis 99:188–194 Fisher JR, Sanchez-Cuevas M-C, Nameth ST, Woods VL, Ellett CW (1997) First report of Cucumber mosaic virus in Eryngium amethystinum, Canna spp., and Aquilegia hybrids in Ohio. Plant Dis 81(11):1331 Jeon YW, Hong JS, Lee SY, Ryu KH, Choi JK (2006) Characterization of an isolate of Cucumber mosaic virus isolated from Canna generalis Bailey. Res Plant Dis 12:298–302 Kumari A, Kumar S, Raj SK (2014) First report of Canna yellow mottle virus on Canna from India. New Dis Rep 29:9 Kumari A, Kaur C, Kumar S, Raj SK, Roy RK, Nautiyal CS (2015) First report of Bean yellow mosaic virus causing a mosaic disease of Canna sp. in India. Plant Dis 99:897 Lockhart B (1988) Occurrence of Canna yellow mottle virus in North America. Acta Hortic 234:69–72 Marino MT, Ragozzino E, Lockhart BEL, Miglino R, Alioto D (2008) First report of Canna yellow mottle virus (CaYMV) in Italy and in the Netherlands. Plant Pathol 57:394 Mitrofanova IV, Zakubanskiy AV, Mitrofanova OV (2018) Viruses infecting main ornamental plants: an overview. Ornamental Hortic 24:95–102 Moini AA, Izadpanah K (2000) New hosts for Tomato spotted wilt virus in Tehran. Iran J Plant Pathol 36:104–105 Momol MT, Lockhart BE, Dankers H, Adkins S (2004) Canna yellow mottle virus detected in canna in Florida. Plant Health Prog. https://doi.org/10.1094/PHP-2004-0809-01-HN Monger WA, Harju V, Skelton A, Seal SE, Mumford RA (2007) Canna yellow streak virus: a new potyvirus associated with severe streaking symptoms in canna. Arch Virol 152:1527–1530 Monger WA, Adams IP, Glover RH, Barrett B (2010) The complete genome sequence of Canna yellow streak virus. Arch Virol 155:1515–1518 Olszewski NE, Lockhart B (2011) Badnavirus. Caulimoviridae. In: The Springer index of Viruses. Springer, New York, pp 263–269. https://doi.org/10.1007/978-0-387-95919-1_40 Pappu HR, Druffel KB, Eastwell KC (2008) Canna yellow mottle virus in Canna spp. in Washington State. Plant Dis 92:1136 Reichel H, Belalcázar S, Munera G, Arevalo E, Narvaez J (1997) First report of Banana streak virus infecting sugarcane and arrowroot in Colombia. Plant Dis 81:552 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Saidi A, Safaeizadeh M (2012) First report of Cucumber mosaic virus on Canna indica in Iran. Aust Plant Dis Notes 7:119–121 Samuitiene M, Navalinskiene M (2008) Occurrence of Cucumber mosaic cucumovirus on ornamental plants in Lithuania. Zemdirbyste-Agric 95:135–143 Tang W, Xu XH, Sun HW, Li F, Gao R, Yang SK, Lu XB, Li XD (2016) First report of Sugarcane mosaic virus infecting Canna spp. in China. Plant Dis 100:2541 Tang W, Yan Z-Y, Zhu T-S, Xu X-J, Li X-D, Tian Y-P (2018) The complete genomic sequence of Sugarcane mosaic virus from Canna spp. in China. Virol J 15:147 Wijayasekara D, Hoyt P, Gimondo A, Thapa A, Jones H, Verchot J (2018) Molecular characterization of two badnavirus genomes associated with Canna yellow mottle disease. Virus Res 243:19–24 Windham A (2008) First report of Canna yellow mottle virus in Tennessee. Ornamental Pest Dis Updat 2:3 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Yamashita S, Natsuaki T, Doi Y, Yora K (1985) Canna yellow mottle virus, a non-enveloped small-bacilliform virus in Canna sp. Ann Phytopath Soc Jpn 51:642–646 Zakubanskiy AV, Mitrofanova IV, Chirkov SN (2017) Molecular characterization of viruses infecting canna in Russia. Eur J Plant Pathol 149:923–931

Capparis spinosa (Caper bush)

367

Cannabis sativa (Hemp) Family: Cannabaceae

Cannabis cryptic virus Taxonomic position Genus: Betapartitivirus

Fiber crop

(CCV)

Family: Partitiviridae

Geographical distribution CCV infection in plants of Cannabis sativa was reported from Germany (Ziegler et al. 2012). Symptoms and host(s) The virus-infected hemp plants do not exhibit any symptoms. Transmission The virus has no known natural vector. The virus is not graft-transmissible. Probably seed transmission is the only known mode of virus spread (Boccardo et al. 1987). Virion properties and genome The virions are isometric, non-enveloped, and 30–43 nm in diameter. The genome is comprised of two molecules of dsRNA with one predicted open reading frame each. RNA1 is 2420 bp (JN196536), and the predicted ORF starts at base pairs 109 and ends at nucleotide 2346. RNA2 is slightly shorter with 2290 bp (JN196537), and the predicted ORF spans base pairs 92 through 2110 (Ziegler et al. 2012; Vainio et al. 2018).

References Boccardo G, Lisa V, Luisoni E, Milne RG (1987) Cryptic plant viruses. Adv Virus Res 32:171–214 Vainio EJ, Chiba S, Ghabrial SA, Maiss E, Roossinck M, Sabanadzovic S, Suzuki N, Xie J, Nibert M, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Partitiviridae. J Gen Virol 99:17–18 Ziegler A, Matouek J, Steger G, Schubert J (2012) Complete sequence of a cryptic virus from hemp (Cannabis sativa). Arch Virol 157:383–385

Capparis spinosa (Caper bush) Family: Capparaceae

Caper latent virus Taxonomic position Genus: Carlavirus

Leafy vegetable

(CapLV)

Family: Betaflexiviridae

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Capparis spinosa (Caper bush)

Geographical distribution CapLV was first reported in plants of Capparis spinosa from Apulia, Southern Italy, by Di Franco and Gallitelli (1985). The virus spreads in Italy (Apulia) (Gallitelli and Di Franco 1987; Tomassoli et al. 2006; Tiberini et al. 2011). Symptoms and host(s) The virus-infected caper bush plants do not exhibit any symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. Experimentally infected plants mostly show chlorotic or necrotic local lesions, systemic mosaic, and leaf malformation. The virus is transmissible by grafting. Virion properties and genome The virions are flexuous filaments about 662 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA 7.4–7.9 kb in size and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Adams et al. 2004; Tiberini et al. 2011). Partial sequences are available (HQ588147, HQ588148).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Capparis spinosa was reported from Italy (Tomassoli et al. 2005, 2006). The virus-infected caper bush plants exhibit symptoms of reddish mottle, vein yellowing, and yellow rings or spots. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Di Franco A, Gallitelli D (1985) Rhabdovirus-like particles in caper leaves with vein yellowing. Phytopathol Mediterr 24:234–236 Gallitelli D, Di Franco A (1987) Characterization of Caper latent virus. J Phytopatol 119:97–105 Tiberini A, Mascia T, Tomassoli L (2011) First nucleotide sequence of a Carlavirus genus infecting caper. Phytopathol Mediterr 50:482–488 Tomassoli L, Zaccaria A, Barba M (2005) Capparis spinosa, a new host of Cucumber mosaic virus in Italy. Plant Pathol 54:263 Tomassoli L, Zaccaria A, Barba M (2006) Virus distribution and incidence on Capparis spinosa L. in Sicilian Islands. In: Proceedings of the 12th congress of Mediterranean Phytopathological Union, 11–15 June Rhodes Island, Greece, pp 125–126

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Capraria spp. Family: Scrophulariaceae

Weed host

Capraria yellow spot virus Taxonomic position Genus: Begomovirus

(CarYSV)

Family: Geminiviridae

Geographical distribution CarYSV infection in plants of Capraria biflora was reported from Mexico (Torres-Herrera et al. unpublished - KC426927). Symptoms and host(s) The virus-infected Capraria plants exhibit yellow spot symptoms on the leaves. Transmission The vector transmission has not been investigated; however, the virus is expected to be transmitted by the whitefly Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm, with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A contains 2624 nt (KC426927 = NC_022007) and DNA-B of 2584 nt (KC426928 = NC_022008) (Briddon 2001; Torres-Herrera et al. unpublished - KC426927; Brown et al. 2015; Zerbini et al. 2017).

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Capsella bursa-pastoris (Shepherd’s purse) Family: Brassicaceae

Beet curly top virus

Weed host

(BCTV)

Synonyms Beet mild curly top virus (BMCTV)

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Taxonomic position Genus: Curtovirus

Capsella bursa-pastoris (Shepherd’s purse)

Family: Geminiviridae

BMCTV infection in plants of Capsella bursa-pastoris was reported from Mexico (Chen et al. 2011). The virus-infected shepherd’s purse plants exhibit symptoms of severely stunted, distorted, and twisted leaves and seedstalks. The virus is transmitted by the leafhopper vector, Circulifer tenellus in a persistent, circulative, non-propagative manner. The virus is not mechanically sap-transmissible. For more details of BCTV, refer to Beta vulgaris.

Beet mild yellowing virus Taxonomic position Genus: Polerovirus

(BMYV)

Family: Luteoviridae

BMYV infection in plants of Capsella bursa-pastoris was reported from France (Hauser et al. 2000). The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of BMYV, refer to Beta vulgaris.

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

BWYV infection in plants of Capsella bursa-pastoris was reported from British Columbia and the USA (Ellis 1992; Falk and Duffus 1984). The virus-infected shepherd’s purse plants exhibit reddening of the lower leaves, interveinal chlorosis, and stunting of plants. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of BWYV, refer to Beta vulgaris.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Capsella bursa-pastoris was reported from New Zealand and Britain (Tomlinson et al. 1970; Fletcher 2001). The virus-infected shepherd’s purse plants exhibit symptoms of mottling of the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

Family: Luteoviridae

(CABYV)

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CABYV infection in plants of Capsella bursa-pastoris was reported from Tunisia (Mnari-Hattab et al. 2009). The virus-infected shepherd’s purse plants exhibit severe yellowing symptoms on older leaves. The virus is transmitted by aphid vectors in a circulative, non-propagative manner, and not transmissible by mechanical sap-inoculation. For more details of CABYV, refer to Cucumis melo.

Pelargonium zonate spot virus Taxonomic position Genus: Anulavirus

(PZSV)

Family: Bromoviridae

PZSV infection in plants of Capsella bursa-pastoris was reported from Spain (Luis-Arteaga and Cambra 2000; Gallitelli et al. 2005). The virus-infected shepherd’s purse plants exhibit symptoms of severe chlorotic and necrotic ringspots, line patterns on leaves, and concentric chlorotic ringspots on stems. Thrips are involved in transmitting this virus by carrying the virus-infected pollen grains on their bodies and introducing the virus into the susceptible hosts while feeding. The virus is mechanically saptransmissible to a wide range of test plants. For more details of PZSV, refer to Pelargonium spp.

Plum pox virus Taxonomic position Genus: Potyvirus

(PPV)

Family: Potyviridae

PPV infection in plants of Capsella bursa-pastoris was reported from Bulgaria (Milusheva and Rankova 2002). The virus-infected shepherd’s purse plants exhibit both local and systemic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of PPV, refer to Prunus domestica.

Potato leafroll virus Taxonomic position Genus: Polerovirus

(PLRV)

Family: Luteoviridae

PLRV infection in plants of Capsella bursa-pastoris was reported from British Columbia (Ellis 1992). The virus-infected shepherd’s purse plants exhibit reddening of the lower leaves, interveinal chlorosis, and stunting of plants. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of PLRV, refer to Solanum tuberosum.

Potato virus Y

(PVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

PVY infection in plants of Capsella bursa-pastoris was reported from New Zealand (Fletcher 2001). The virus-infected shepherd’s purse plants exhibit mosaic symptoms. The virus is transmitted by aphid

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Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts and by grafting. For more details of PVY, refer to Solanum tuberosum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Capsella bursa-pastoris was reported from Hawaii and British Columbia (Cho et al. 1986; Bitterlich and MacDonald 1993). The virus-infected shepherd’s purse plants exhibit yellow spot and ringspot symptoms. The virus is transmitted by thrips vectors in a persistentpropagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Bitterlich I, MacDonald LS (1993) The prevalence of Tomato spotted wilt virus in weeds and crops in southwestern British Columbia. Can Plant Dis Surv 73:137–142 Chen L-F, Vivoda E, Gilbertson RL (2011) Genetic diversity in curtoviruses: a highly divergent strain of Beet mild curly top virus associated with an outbreak of curly top disease in pepper in Mexico. Arch Virol 156(4):547–555 Cho JJ, Mau RFL, Gonsalves D, Mitchell WC (1986) Reservoir weed hosts of Tomato spotted wilt virus. Plant Dis 70:1014–1017 Ellis PJ (1992) Weed hosts of Beet western yellows virus and Potato leafroll virus in British Columbia. Plant Dis 76:1137–1139 Falk BW, Duffus JE (1984) Identification of small single- and double-stranded RNA associated with severe symptoms in Beet western yellows virus – infected Capsella bursa-pastoris. Phytopathology 74:1224–1229 Fletcher D (2001) New hosts of Alfalfa mosaic virus, Cucumber mosaic virus, Potato virus Y, Soybean dwarf virus, and Tomato spotted wilt virus in New Zealand. N Z J Crop Hortic Sci 29:213–217 Gallitelli D, Finetti-Sialer M, Martelli GP (2005) Anulavirus, a proposed new genus of plant viruses in the family Bromoviridae. Arch Virol 150:407–411 Hauser S, Stevens M, Mougel C, Smith HG, Fritsch C, Herrback E, Lemaire O (2000) Biological, serological and molecular variability suggest three distinct polerovirus species infecting beet or rape. Phytopathology 90:460–466 Luis-Arteaga M, Cambra M (2000) First report of natural infection of greenhouse-grown tomato and weed species by Pelargonium zonate spot virus in Spain. Plant Dis 84:807 Milusheva S, Rankova Z (2002) Plum pox potyvirus detection in weed species under field conditions. Acta Hortic 577:283–287 Mnari-Hattab M, Gauthier N, Zouba A (2009) Biological and molecular characterization of the Cucurbit aphid-borne yellows virus affecting cucurbits in Tunisia. Plant Dis 93:1065–1072 Tomlinson JA, Carter AL, Dale WT, Simpson CJ (1970) Weed plants as sources of Cucumber mosaic virus. Ann Appl Biol 66:11–16

Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper) Family: Solanaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Spice crop

(AMV)

Family: Bromoviridae

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AMV infection in plants of Capsicum annuum was widely distributed in Saudi Arabia, Mediterranean basin, Zambia, Turkey, the Czech Republic, Ecuador, Yemen, Serbia, and the USA (Alhubaishi et al. 1987; Hamm et al. 1995; Ndunguru and Kapooria 1999; Marchoux et al. 2003; Petrovic et al. 2010; Abdalla and Ali 2012; Svoboda and Svobodova-Leisova 2012; Al-Abrahaim 2013; Colimba et al. 2016; Al-Shahwan et al. 2017). The virus-infected pepper plants develop local chlorotic and necrotic rings on the leaves followed by systemic foliar symptoms on leaves with distinct yellow or whitish mosaic symptoms. Some AMV isolates cause severe necrosis in certain pepper cultivars. Relative to uninfected plants, fruit may be stunted and misshapen. The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Alstroemeria necrotic streak virus

(ANSV)

Taxonomic position ANSV is a tentative member of the genus Orthotospovirus and family Tospoviridae ANSV infection in plants of Capsicum annuum was reported from Colombia (Olaya et al. 2017). The virus-infected bell pepper plants exhibit necrotic lesions, necrotic rings, veinal necrosis, and stem necrosis symptoms. The virus is transmitted by a thrips vector Frankliniella occidentalis in a persistent propagative manner. For more details of ANSV, refer to Alstroemeria spp.

Andean potato mottle virus Taxonomic position Genus: Comovirus

(APMoV)

Family: Secoviridae

APMoV infection in plants of Capsicum frutescens was reported from Central America (Valverde et al. 1995). The virus-infected bell pepper plants exhibit mosaic and yellow mottling symptoms. The virus is transmitted by the cucumber beetle vector, Diabrotica balteata, in a non-persistent manner. The virus is mechanically sap-transmissible and also by contact between plants. For more details of APMoV, refer to Solanum tuberosum.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Capsicum annuum was reported from Saudi Arabia (Al-Shahwan et al. 2017). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Beet curly top virus

(BCTV)

Synonyms Beet mild curly top virus (BMCTV); Beet severe curly top virus (BSCTV); Pepper yellow dwarf virus (PepYDV); Texas pepper geminivirus

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Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

Taxonomic position Genus: Curtovirus

Family: Geminiviridae

BMCTV was reported in plants of Capsicum annuum in the USA, Mexico, Costa Rica, and Iran (Stenger et al. 1990; Hamm et al. 1995; Guzman et al. 1996; Lotrakul et al. 2000; Velasquez-Valle et al. 2008; Lam et al. 2009; Chen et al. 2011, 2017; Robles-Hernandez et al. 2011; Mauricio-Castillo et al. 2017). The Early in the season virus-infected pepper plants are chlorotic and stunted with thickened, elongated leaves. From mid- to late season, the affected plants show severe yellowing, upwardly rolled, small leaves, and a few deformed fruits. The virus is transmitted by the leafhopper vector, Circulifer tenellus in a persistent circulative, non-propagative manner. The virus is not mechanically saptransmissible. For more details of BCTV, refer to Beta vulgaris.

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

BWYV infection in plants of Capsicum frutescens was reported from China (Yuan et al. 2015). The virus-infected pepper plants show mosaic and crinkling of the young leaves. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of BWYV, refer to Beta vulgaris.

Bell pepper alphaendornavirus Taxonomic position Genus: Alphaendornavirus

(BPEV)

Family: Endornaviridae

Geographical distribution BPEV infection in plants of Capsicum annuum was reported from the USA, Japan, and Canada (Valverde et al. 1990; Valverde and Gutierrez 2007; Okada et al. 2011; Chen et al. 2015). Symptoms and host(s) The virus-infected pepper cultivars does not produce any symptoms. The virus was also detected in other Capsicum species (Capsicum annuum, C. baccatum, C. Chinense, and C. frutescens). Transmission The virus is transmitted through seed via both ova and pollen. No horizontal spread has been observed in the field and no potential vectors have been identified. The virus is not mechanically sap-transmissible and also not transmissible by grafting (Valverde and Gutierrez 2007). Virion properties and genome The virus does not have a gene for a capsid protein; therefore, no virion is formed. The virus has a linear dsRNA of 14,727 bp as its genome, which contains a single, long ORF (starting at nt 226 and ending at nt 14,670) encoding a polyprotein of 4815 amino acids (JN019858, AB597230). The virus contains a nick in the positive-strand RNA molecule. The virus has conserved motifs of viral methyltransferase,

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helicase 1, UDP-glycosyltransferase, and RNA-dependent RNA polymerase (RdRp) (Okada et al. 2011; Sela et al. 2012).

Bell pepper mottle virus Taxonomic position Genus: Tobamovirus

(BPMoV)

C Family: Virgaviridae

Geographic distribution BPMoV infection in plants of Capsicum annuum was originally reported from Argentina and later was also found in the Netherlands (Wetter et al. 1987; Green and Kim 1991). The virus spreads in Germany, Netherlands, and Argentina. Symptoms and host(s) The virus-infected pepper plants show mottling symptoms. Transmission Transmission occurs without the help of vectors. The virus is mechanically sap-transmissible and also through contact between plants. Virion properties and genome The virions are non-enveloped, rigid helical rods with a helical symmetry, about 18 nm in diameter, and 300 nm in length. The genome is a monopartite, linear, positive-sense ssRNA of 6375 nt (DQ355023 = NC_009642). The 30 -terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G50 pppG) (Rhie et al. 2007; Zaitlin 2011; Adams et al. 2017).

Bhendi yellow vein mosaic virus Taxonomic position Genus: Begomovirus

(BYVMV)

Family: Geminiviridae

BYVMV infection in plants of Capsicum annuum was reported from India (Singh et al. 2015). The virus-infected pepper plants exhibit symptoms of malformation, leaf curling, vein-clearing, and stunting. The virus is transmitted by a whitefly vector; Bemisia tabaci in a circulative non-propagative manner. The virus is not mechanically sap-transmissible. For more details of BYVMV, refer to Abelmoschus esculentus (Syn.) Hibiscus esculentus.

Broad bean wilt virus 1 Taxonomic position Genus: Fabavirus

(BBWV-1)

Family: Secoviridae

BBWV-1 infection in plants of Capsicum annuum was reported from Slovenia, the Czech Republic, and Spain (Rubio et al. 2002; Mehle et al. 2008; Svoboda and Svobodova-Leisova 2012). The virus-infected pepper plants exhibit chlorotic blotching symptoms. The virus is transmitted by

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Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BBWV-1, refer to Vicia faba.

Broad bean wilt virus 2 Taxonomic position Genus: Fabavirus

(BBWV-2)

Family: Secoviridae

BBWV-2 infection in plants of Capsicum annuum was reported from the Czech Republic and Korea (Lee et al. 2000; Choi et al. 2005; Kwak et al. 2013; Svoboda and Leisova-Svobodova 2013). The virusinfected pepper plants exhibit systemic mottling symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BBWV-2, refer to Vicia faba.

Capsicum chlorosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(CaCV)

Family: Tospoviridae

Geographical distribution CaCV infection in plants of Capsicum annuum was reported from India, Thailand, and Australia (McMichael et al. 2002; Jones and Sharman 2005; Persley et al. 2006; Krishnareddy et al. 2008; Kunkalikar et al. 2010, 2011; Bhat et al. 2014; Sharma and Kulshrestha 2016; Basavaraj et al. 2017). Symptoms and host(s) Young leaves of CaCV-infected pepper plants show marginal and interveinal chlorosis and often become narrow and curled. Older leaves are chlorotic and may develop ringspot and line pattern symptoms. Infected plants are often stunted with small, distorted fruit that develop necrotic lesions and scarring (McMichael et al. 2002; Krishnareddy et al. 2008). The natural host range of this virus includes multiple vegetable (tomato, chilli, capsicum), legume (peanut) and some ornamental plants. Transmission The virus is transmitted by tomato thrips (F. schulzei) and melon thrips (Thrips palmi). Thrips species such as plague thrips (T. imaginis) and greenhouse thrips (Heliothrips haemmorhoidalis) do not transmit the virus. The virus is mechanically sap-transmissible to capsicum, tomato, peanut, and several weed species (Jones and Sharman 2005). Virion properties and genome The virions consist of a quasi-spherical capsid particle 80–120 nm in diameter. The genome consists of three linear single-stranded RNAs (L: 8912 nt (DQ256124); M: 4823 nt (DQ256125); S: 3477 nt (DQ256123)) (Manyam and Byadgi 2014; Gamage et al. 2015; Huang et al. 2017a).

Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

Chickpea chlorotic dwarf virus Taxonomic position Genus: Mastrevirus

377

(CpCDV)

Family: Geminiviridae

CpCDV infection in plants of Capsicum annuum was reported from Oman and India (Akhtar et al. 2014; Byun et al. 2014). The virus-infected pepper plants exhibit symptoms of upward leaf curling leading to cupping and stunting. The virus is transmitted by leafhopper vectors in a persistent, circulative, and non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of CpCDV, refer to Cicer arietinum.

Chilli leaf curl Ahmedabad virus Taxonomic position Genus: Begomovirus

(ChiLCAV)

Family: Geminiviridae

Geographical distribution ChiLCAV infection in plants of Capsicum annuum was reported from Ahmedabad, India (Bhatt et al. 2016). Symptoms and host(s) The virus-infected chilli plants exhibit leaf curl symptoms. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, nonpropagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2744 nt (KM880103 = NC_028046) (Brown et al. 2015; Bhatt et al. 2016; Zerbini et al. 2017).

Chilli leaf curl Bhavanisagar virus Taxonomic position Genus: Begomovirus

(ChiLCBV)

Family: Geminiviridae

Geographical distribution ChiLCBV infection in plants of Capsicum annum was reported from Tamil Nadu, India (Parvathy Krishnan et al. 2010). Symptoms and host(s) The virus-infected chilli plants exhibit severe curling and reduced leaf size. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, nonpropagative manner.

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Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2785 nt (HM992939) (Parvathy Krishnan et al. 2010; Brown et al. 2015; Zerbini et al. 2017).

Chilli leaf curl Gonda virus Taxonomic position Genus: Begomovirus

(ChiLCGV)

Family: Geminiviridae

Geographical distribution ChiLCGV infection in plants of Capsicum annum was reported from Gonda, India (Khan and Khan 2017). Symptoms and host(s) The virus-infected chilli plants show leaf curl symptoms. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, nonpropagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2760 nt (KJ957157) (Brown et al. 2015; Khan and Khan 2017; Zerbini et al. 2017).

Chilli leaf curl India virus Taxonomic position Genus: Begomovirus

(ChiLCINV)

Family: Geminiviridae

Geographical distribution ChiLCINV infection in plants of Capsicum annuum was reported from India (Kumar et al. unpublished FM877858). Symptoms and host(s) The virus-infected pepper plants exhibit severe leaf curl and puckering symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2755 nt (FM877858) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

Chilli leaf curl Kanpur virus Taxonomic position Genus: Begomovirus

379

(ChiLCKaV)

Family: Geminiviridae

Geographical distribution ChiLCKaV infection in plants of Capsicum annuum was reported from India (Singh et al. unpublished HM007106. Symptoms and host(s) The virus-infected pepper plants exhibit leaf curling and puckering symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2754 nt (HM007106) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Chilli leaf curl Palampur virus

(ChiLCPaV)

Taxonomic position ChiLCPaV is a tentative member of the genus Begomovirus and family Geminiviridae Geographical distribution ChiLCPaV infection in plants of Capsicum frutescens was reported from Palampur, India (Kumar et al. 2011). Symptoms and host(s) The virus-infected bell pepper plants exhibit leaf curl and puckering symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The virions have geminate morphology measuring 18  30 nm. The genome is circular single-stranded DNA of 2775 nt. A betasatellite DNA molecule is associated with the virus, and consists of 1376 nt (Kumar et al. 2011).

Chilli leaf curl Sri Lanka virus Taxonomic position Genus: Begomovirus

(ChiLCSLV)

Family: Geminiviridae

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Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

Geographical distribution ChiLCSLV infection in plants of Capsicum annuum was known to occur in Sri Lanka (Senanayake et al. 2013, 2015). Symptoms and host(s) The virus-infected chilli plants exhibit symptoms of reduced leaf size in younger leaves and boatshaped leaves with bushy appearance, margins rolled upwards, and necrosis at tip, or reduced leaf size, with vein clearing followed by leaf curl. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. The virus is readily transmitted by Bemisia tabaci to chilli, tomato, and tobacco, where vein-clearing developed followed by leaf curl (Senanayake et al. 2013). Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2754 nt (JN555601) (Senanayake et al. 2013, 2015; Brown et al. 2015; Zerbini et al. 2017). Chilli leaf curl Sri Lanka betasatellite DNA molecule is associated with ChiLCSLV, and consists of 1371 nt (JN638445) (Zhou 2013).

Chilli leaf curl Vellanad virus Taxonomic position Genus: Begomovirus

(ChiLCVV)

Family: Geminiviridae

Geographical distribution ChiLCVV infection in plants of Capsicum annuum was reported from Vellanad district of Kerala (Vinoth Kumar et al. 2012). Symptoms and host(s) The virus-infected pepper plants exhibit symptoms of typical upward leaf curling, crinkling, puckering, and reduction of leaf area along with stunting of whole plants. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm, with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2,788 nt (HM007121) with seven predicted ORFs (AV1, AV2, AC1, AC2, AC3, AC4, and AC5). Association of two betasatellites 1355 nt (JN663876) and 1372 nt (JN663877) each containing a single ORF C1 is reported (Briddon 2001; Vinoth Kumar et al. 2012; Brown et al. 2015; Zerbini et al. 2017).

Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

Chilli leaf curl virus Taxonomic position Genus: Begomovirus

381

(ChiLCV)

Family: Geminiviridae

Geographical distribution ChiLCV infection in plants of Capsicum annuum was known to occur in India, Oman, and Pakistan (Shih et al. 2003; Senanayake et al. 2007, 2012; Khan et al. 2013; Kaur et al. 2015). Some isolates of the virus have previously been known as Pepper leaf curl Pakistan virus, Chilli leaf curl Multan virus, and Chilli leaf curl Salem virus. Symptoms and host(s) The virus-infected pepper plants exhibit yellowing, upward curling, puckering, and a reduced size. Severely affected plants are stunted and produced no fruit. The virus has been isolated from chilli (Capsicum annuum), eggplant (Solanum melongena), green amaranth (Amaranthus viridis), kenaf (Hibiscus cannabinus), papaya (Carica papaya), Phaseolus aureus, potato (Solanum tuberosum), and tomato (Solanum lycopersicum). Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative and non-propagative manner (Senanayake et al. 2012). A single whitefly was capable of transmitting ChiLCV, and eight or more whiteflies per plant resulted in 100% transmission. The minimum acquisition access period (AAP) and inoculation access period (IAP) were 180 and 60 min, respectively. The virus persisted in whiteflies for up to 5 days postacquisition (Senanayake et al. 2012). Experimentally, the virus has been shown to infect tomato (Solanum lycopersicum), Nicotiana tabacum, N. benthamiana and papaya (Carica papaya). Virion properties and genome For some isolates, the presence of geminate particles, typical of geminiviruses, has been shown in infected plants (Senanayake et al. 2012). Nevertheless, the fine structure of ChiLCV particles has not been investigated. The genome of ChiLCV consists of a single, circular single-stranded DNA of ~2754 nt (AF336806 = NC_004628) (Briddon 2001; Senanayake et al. 2012; Brown et al. 2015). Chilli leaf curl betasatellite DNA molecule is associated with ChiLCV, and consists of 1387 nt (AJ316032) (Briddon et al. 2003; Zhou 2013; Zerbini et al. 2017). The characterized genomes of ChiLCV encode the six genes typically encoded by monopartite begomoviruses. The expression and function of the genes have not been investigated.

Chilli ringspot virus Taxonomic position Genus: Potyvirus

(ChiRSV)

Family: Potyviridae

ChiRSV infection in plants of Capsicum annuum var. sanam was reported from Pakistan and Laos (Ahmad et al. 2017; Chittarath et al. 2017). The virus-infected chilli pepper plants exhibit symptoms of mottling, interveinal chlorosis, and dark green vein-banding symptoms. The virus is transmitted by

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aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ChiRSV, refer to Capsicum chinense.

Chilli veinal mottle virus

(ChiVMV)

Synonyms Indian pepper mottle virus; Indian pepper vein banding virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution ChiVMV infection in plants of Capsicum annuum was first reported from Malaysia (Burnett 1947). The virus spreads in Korea, Philippines, Taiwan, China, India, Japan, Nigeria, South Africa, Brazil, Pakistan, Italy, and Thailand (Sulaiman and Gim 1981; Ravi 1991; Ravi et al. 1997; Joseph 1999; Shah et al. 2008, 2009; Tsai et al. 2008; Tiberini et al. 2017). Symptoms and host(s) The virus-infected pepper plants exhibit symptoms of leaf mottle and dark green vein-banding, veinclearing, deformation, chlorotic streaks and leaves may be small and distorted. If plants are infected when young, they may become stunted, with dark green streaks on their stems and branches and malformed fruits. The host range of the virus is restricted to only the family Solanaceae (Shah et al. 2008). Transmission The virus is transmitted by aphid vectors, Aphis craccivora, A. gossypii, A. spiraecola, Myzus persicae, Toxoptera citricidus, Hysteroneura setarieae, Rhopalosiphum maidis in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and by grafting and not by seed (Bidari and Reddy 1990; Moi 1992; Moury et al. 2005). Virion properties and genome The virions are non-enveloped, flexuous filaments 648–759 nm long and 13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9711 nt (AJ237843 = NC_005778) (Green 1999; Joseph 1999; Joseph and Savithri 1999; Anindya et al. 2004; Hwang et al. 2009; Ha et al. 2013; Sharma et al. 2014; Wylie et al. 2017).

Cotton leaf curl Kokhran virus Taxonomic position Genus: Begomovirus

(CLCuKoV)

Family: Geminiviridae

CLCuKoV infection in plants of Capsicum annuum was reported from Pakistan (Yasmin et al. 2017). The virus-infected chilli pepper plants exhibit leaf curl symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. The virus is not transmissible by mechanical inoculation or by contact between plants. For more details of CLCuKoV, refer to Gossypium spp.

Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

Cowpea aphid-borne mosaic virus Taxonomic position Genus: Potyvirus

383

(CABMV)

Family: Potyviridae

CABMV infection in plants of Capsicum annuum was reported from Nigeria (Aliyu 2014). The virus is transmitted by a number of aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of CABMV, refer to Vigna unguiculata.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Capsicum annuum distributed all over the world including Asia, Egypt, Italy, Korea, Spain, California, Zambia, Venezuela, Serbia, and Argentina (Simons 1957; Anjaneyulu and Apparao 1967; Lockhart and Fisher 1976; Debrot 1980; Nagaraju and Reddy 1982; Tobias et al. 1982; Florini and Zitter 1987; Abdel-Salam et al. 1989; Subasic et al. 1990; George et al. 1993; Atencio et al. 1997; Lapidot et al. 1997; Ndunguru and Kapooria 1999; Ben Chaim et al. 2001; Kim et al. 2002; Rodriguez-Alvarado et al. 2002; Frangioni et al. 2003; Rao and Lan 2003; Bhadramurthy et al. 2009; Petrovic et al. 2010; Choi et al. 2011; Sharma and Patiyal 2011; Arogundade et al. 2012; Kapoor 2012; Biswas et al. 2013, 2016; Myti et al. 2014; Olawale et al. 2015; Ashwathappa et al. 2016). The virus commonly infects chilli pepper and causes different type of symptoms depending on the strain of CMV, genotype of plant and stage of infection. The most common symptom in chilli is stunting of plants, with pale and leathery foliage. The leaves sometimes show chlorotic mosaic pattern, necrotic specks, or ringspots. The shape of leaf may become narrow and leaves may drop prematurely. Mosaic mottling on leaves may also be noticed. Fruit have a wrinkled, bumpy appearance, uneven color, and ripening with dark spots sometimes developing. The virus is transmitted by more than 86 spp. of aphid vectors Aphis gossypii, A. craccivora, Acyrthosiphon pisum, and Myzus persicae which are the major vectors. The virus-vector relationship is the non-persistent manner. The virus is mechanically sap-transmissible. The virus is transmissible through seed of Capsicum annuum up to 53–83% (Ali and Kobayashi 2010). For more details of CMV, refer to Cucumis sativus.

Dahlia mosaic virus Taxonomic position Genus: Caulimovirus

(DMV)

Family: Caulimoviridae

DMV infection in plants of Capsicum annuum was reported from China (Zhang et al. 2015b). The virus-infected pepper plants exhibit leaf chlorosis and mild upward leaf curl symptoms. The virus is transmitted by aphid vectors in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of DMV, refer to Dahlia spp.

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Eggplant mottled dwarf nucleorhabdovirus Taxonomic position Genus: Nucleorhabdovirus

(EMDV)

Family: Rhabdoviridae

EMDV infection in plants of Capsicum annuum was reported from Bulgaria (Roggero et al. 1995; Kostova et al. 2001). The virus-infected pepper plants exhibit symptoms of severe vein-clearing and small yellow spots or yellow mosaic on young leaves. The virus is transmitted by a leafhopper vector, Agallia vorobjevi in a persistent, propagative manner. The virus is mechanically saptransmissible to solanaceous hosts and also by grafting. For more details of EMDV, refer to Solanum melongena.

Groundnut bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(GBNV)

Family: Tospoviridae

GBNV infection in plants of Capsicum annuum was reported from India (Kunkalikar et al. 2011; Bhat et al. 2014; Sharma and Patiyal 2011; Pavithra et al. 2016; Sharma and Kulshrestha 2016). The virusinfected pepper plants exhibit chlorotic ringspots on leaves and necrosis on stems. The virus is transmitted by a thrips vector in a persistent propagative manner, and also by mechanical sapinoculation. For more details of GBNV, refer to Arachis hypogaea.

Groundnut ringspot orthotospovirus Taxonomic position Genus: Orthotospovirus

(GRSV)

Family: Tospoviridae

GRSV infection in plants of Capsicum annuum was reported from Brazil, South Africa, South America, and the United States (Colariccio et al. 2001; Webster et al. 2015). The virus-infected pepper plants exhibit symptoms of chlorotic and necrotic spots on leaves, inward rolling of leaves, and general plant stunting. Fruits of infected plants are generally discolored, often deformed, and with chlorotic or necrotic ring patterns. The virus is transmitted by thrips vectors, F. occidentalis and F. schultzei, in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of GRSV, refer to Arachis hypogaea.

Groundnut yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(GYSV)

Family: Tospoviridae

GYSV infection in plants of Capsicum annuum was reported from China (Ding et al. 2007). The virusinfected pepper plants exhibit severe necrotic ringspot symptoms. The virus is transmitted by thrips vectors in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of GYSV, refer to Arachis hypogaea.

Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

Hot pepper alphaendornavirus Taxonomic position Genus: Alphaendornavirus

385

(HpEV)

Family: Endornaviridae

Geographical distribution HpEV infection in plants of Capsicum annuum was reported from Korea (Lim et al. 2015). Symptoms and host(s) The virus-infected pepper plants do not exhibit any external symptoms. Transmission The virus is transmitted through seed via both ova and pollen. No horizontal spread has been observed in the field and no potential vectors have been identified. The virus is not mechanically sap-transmissible. Virion properties and genome The virus does not have a gene for a capsid protein; therefore, no virion is formed. The genome is double-stranded RNA of 14,729 bp, including 12 cytosines (Cs) at the 30 end (KR080326) (Lim et al. 2015). The putative single, large open reading frame encodes a 4,884-amino-acid-long polyprotein that contains four putative functional domains: a viral methyltransferase, a viral RNA helicase, a glycosyltransferase, and an RNA-dependent RNA polymerase (Lim et al. 2015).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Capsicum annuum was reported from Europe, Slovenia, France, Spain, Israel, Italy, China, and the USA (Roggero et al. 1999; Vicchi et al. 1999; Naidu et al. 2005; Gonzalez-Pacheco and Silva-Rosales 2013; Chen et al. 2016). Symptoms incited by INSVon pepper leaves show large water soaked necrotic areas. With time, these leaves show veinal necrosis and mild chlorosis. The infected leaves eventually drop from the infected plants. Early infection with INSV leads to severe stunting of pepper plants and necrosis of stem and petioles. Fruits from infected plants show uneven ripening and concentric rings, thereby affecting the quality of the fruit. The virus is vectored by two different species of thrips, Frankliniella occidentalis and F. fusca in a persistent-propagative manner. The virus is also mechanically sap-transmissible to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Melon aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(MABYV)

Family: Luteoviridae

MABYV infection in plants of Capsicum annuum was reported from China (Wang et al. 2017). The virus-infected pepper plants exhibit yellowing and thickening of the leaves. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. For more details of MABYV, refer to Cucumis melo.

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Melon chlorotic leaf curl virus

(MCLCuV)

Synonyms Squash yellow mild mottle virus (SYMMoV) Taxonomic position Genus: Begomovirus

Family: Geminiviridae

SYMMoV infection in plants of Capsicum annuum was reported from Costa Rica (Castro et al. 2013). The virus is transmitted by a whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of MCLCuV, refer to Cucumis melo.

Merremia mosaic virus Taxonomic position Genus: Begomovirus

(MerMV)

Family: Geminiviridae

MerMV infection in plants of Capsicum annuum was reported from Belize (McLaughlin et al. 2008). The virus-infected pepper plants exhibit mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of MerMV, refer to Merremia spp.

Moroccan pepper virus

(MPV)

Synonyms Lettuce necrotic stunt virus (LNSV) Taxonomic position Genus: Tombusvirus

Family: Tombusviridae

Geographical distribution MPV infection in plants of Capsicum annuum was first reported from Morocco by Fischer and Lockhart (1977). The virus spreads in Morocco (Wintermantel and Hladky 2013). Symptoms and host(s) The virus-infected pepper plants exhibit leaf deformation, leaf and flower abscission, and severe systemic necrosis in pepper plants. The natural hosts of this virus are tomato, pelargonium, lisianthus (Eustoma grandiflorum), and Datura stramonium. Transmission There is no vector is known for this virus. The virus is transmitted by mechanical sap-inoculation to 3–9 families. Experimentally infected plants mostly show necrotic local lesions, systemic necrosis, and/or stunting. The virus is highly contagious and it persists in soil, water, and plant debris. The virus is transmissible by grafting.

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Virion properties and genome The virions are isometric and icosahedral (T = 3), non-enveloped and about 32–35 nm in diameter. The genome is a monopartite, linear, positive-sense, single-stranded RNA of 4,772 nt (JX197070, JX197071 and JX182425). The genome contains five ORFs: the ORF1 encodes a 33-kDa protein and the ORF2 encodes a read-through protein of 92-kDa. Both the P33 and P92 proteins are associated with replication. The ORF3 encodes the 41-kDa virus coat protein, and the ORF4 and ORF5 encode the P22 and P19 proteins, respectively (White 2011; Wintermantel and Hladky 2013).

Obuda pepper virus Taxonomic position Genus: Tobamovirus

(ObPV)

Family: Virgaviridae

Geographical distribution ObPV infection in plants of Capsicum annuum was reported from Hungary (Csillery et al. 1983). Symptoms and host(s) The virus infects peppers systemically. Transmission No insect vector is known for this virus. The virus is transmitted through mechanical sap-inoculation, and also by contact between plants. Virion properties and genome The virions are non-enveloped, rigid, helical rods with a helical symmetry, about 18 nm in diameter, and 300–310 nm in length. The genome is a monopartite, linear, positive-sense ssRNA of 6507 nt (D13438 = NC_003852; L11665). The 30 -terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G50 pppG) (Zaitlin 2011; Adams et al. 2017).

Papaya leaf curl virus Taxonomic position Genus: Begomovirus

(PaLCuV)

Family: Geminiviridae

PaLCuV infection in plants of Capsicum annuum was reported from Pakistan (Yasmin et al. 2017). The virus-infected chilli pepper plants exhibit leaf curl symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. The virus is not mechanically saptransmissible. For more details of PaLCuV, refer to Carica papaya.

Paprika mild mottle virus Taxonomic position Genus: Tobamovirus

(PaMMV)

Family: Virgaviridae

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Geographical distribution PaMMV infection in plants of Capsicum annuum was first recorded in the Netherlands during the late 1970s (Rast 1979) and subsequently found in Eurasian region including Bulgaria, Japan (Hamada et al. 2003).

Symptoms and host(s) The virus-infected pepper plants show mild mottle mosaic symptoms. The virus has a narrow host range (Hamada et al. 2003; Yordanova and Stoimenova 2008–2009).

Transmission There is no known vector for this virus. The virus is transmitted by mechanical sap-inoculation to species in less than three families. Experimentally infected plants mostly show small chlorotic or necrotic local lesions, mosaic, and stunting. The virus is also transmissible by contact between plants.

Virion properties and genome The virions are rod shaped 300 nm long and 18 nm wide (Yordanova and Stoimenova 2008–2009). The genome consists of a single molecule of positive-sense ssRNA of 6524 nt (AB089381 = NC_004106). The 30 -terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G50 pppG) (Hamada et al. 2003; Matsumoto et al. 2009; Zaitlin 2011; Adams et al. 2017).

Parietaria mottle virus Taxonomic position Genus: Ilarvirus

(PMoV)

Family: Bromoviridae

PMoV infection in plants of Capsicum annuum was reported from France, Spain, and Italy (Janssen et al. 2005; Parrella et al. 2016). The virus-infected pepper plants exhibit a mosaic and necrosis of the apical leaves, which progresses to the stem, causing stem necrosis and also top necrosis, affected pepper fruits show brown patches and corky rings on the surface. The virus is transmitted by the thrips vectors, presents in/on pollen, and enters into the host through the injuries caused by thrips while feeding. The virus is mechanically sap-transmissible and also through pollen and seed (Aramburu et al. 2010). For more details of PMoV, refer to Parietaria officinalis.

Pedilanthus leaf curl virus Taxonomic position Genus: Begomovirus

(PeLCV)

Family: Geminiviridae

PeLCV infection in plants of Capsicum annuum was reported from Pakistan (Yasmin et al. 2017). The virus-infected chilli pepper plants exhibit leaf curl symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of PeLCV, refer to Pedilanthus tithymaloides.

Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

Pelargonium zonate spot virus Taxonomic position Genus: Anulavirus

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(PZSV)

Family: Bromoviridae

PZSV infection in plants of Capsicum annuum was reported from Spain (Escriu et al. 2009). The virusinfected pepper plants exhibit severe chlorotic ringspots and line pattern symptoms. Thrips are involved in transmitting this virus by carrying the virus-infected pollen grains on their bodies and introducing the virus into the susceptible hosts while feeding. The virus is mechanically sap-transmissible to a wide range of test plants. For more details of PZSV, refer to Pelargonium spp.

Pepper chat fruit viroid Taxonomic position Genus: Pospiviroid

(PCFVd)

Family: Pospiviroidae

Geographical distribution PCFVd was first reported in plants of Capsicum annuum from the Netherlands (Verhoeven et al. 2009). The viroid spreads in the Spain, the Netherlands, and Canada (Verhoeven et al. 2011). Symptoms and host(s) The viroid-infected pepper plants exhibit reduced fruit size with plant growth slightly reduced (Verhoeven et al. 2011). The experimental host range of this viroid is Datura metel, D. stramonium, Nicotiana benthamiana, N. glutinosa, N. rustica, N. tabacum, Petunia  hybrida, Physalis alkekengi, Solanum lycopersicum, S. melongena, S. muricatum, and S. tuberosum (Yanagisawa and Matsushita 2017) Transmission The viroid is transmitted by mechanical sap-inoculation to several solanaceous plant species inducing vein-necrosis and reduced fruit and tuber size in tomato and potato, respectively. Seed transmission is also reported in capsicum to the extent of 19% (Verhoeven et al. 2009). Virion properties and genome The genome consists of a single-stranded circular RNA of 348 nt, and with minor modifications, it has the central conserved and the terminal conserved regions (FJ409044) (Gora-Sochacka 2004; Verhoeven et al. 2009, 2011; Giguere et al. 2014).

Pepper chlorotic spot virus Taxonomic position The virus is a tentative member of the genus Orthotospovirus and family Tospoviridae Geographical distribution The virus infection in plants of Capsicum annuum was reported from Taiwan and China (Cheng et al. 2014; Huang et al. 2017b; Zheng et al. 2017).

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Symptoms and host(s) The virus-infected pepper plants exhibit symptoms of mottling and deformation on leaves and fruits. Transmission The virus is transmitted by thrips vector in a persistent, propagative manner. Virion properties and genome The particles are 80–120 nm in diameter. The genome consists of one negative and two ambisense single-stranded RNAs. The S, M, and L RNAs of PCSV-14YV733 are 3310 nt, 4711 nt, and 8913 nt, respectively (Huang et al. 2017b; Zheng et al. 2017).

Pepper cryptic virus 1 Taxonomic position Genus: Deltapartitivirus

(PCV1)

Family: Partitiviridae

Geographical distribution PCV1 infection in plants of Capsicum annuum was reported from the USA (Valverde and Gutierrez 2008; Sabanadzovic and Valverde 2011). Symptoms and host(s) The virus-infected pepper plants do not exhibit any symptoms. Transmission The virus is transmissible through pollen/seed but cannot be transmissible by grafting or mechanical sap-inoculation, and does not have any known natural vectors. Virion properties and genome The virions are isometric, non-enveloped about 30 nm in diameter. The genome consists of two molecules of linear double-stranded RNA. RNA1 is 1563 bp (JN117276) and RNA2 is 1512 bp (JN117277) (Sabanadzovic and Valverde 2011; Vainio et al. 2018).

Pepper cryptic virus 2 Taxonomic position Genus: Deltapartitivirus

(PCV2)

Family: Partitiviridae

Geographical distribution PCV2 infection in plants of Capsicum annuum was reported from the USA, China, and India (Sabanadzovic and Valverde 2011; Saritha et al. 2016; Sun et al. 2017). Symptoms and host(s) The virus-infected pepper plants do not exhibit any symptoms.

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Transmission The virus is transmissible through pollen/seed but cannot be transmissible by grafting or mechanical sap-inoculation, and does not have any known natural vectors. Virion properties and genome The virions are isometric about 30 nm in diameter. The genome consists of two molecules of linear double-stranded RNA. RNA1 consists of 1609 bp (JN117278) and RNA2 is 1525 bp (JN117279) (Sabanadzovic and Valverde 2011; Vainio et al. 2018).

Pepper golden mosaic virus

(PepGMV)

Synonyms Pepper Texas bigeminivirus, Serrano golden mosaic virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution PepGMV infection was first reported in plants of Capsicum annuum cv. Serrano, from Sinaloa, Mexico, by Brown and Poulos (1990). The virus spreads in the USA, Mexico, Costa Rica, Guatemala, Nicaragua, and Cuba (De La Torre et al. 1998; Lotrakul et al. 2000; Idris et al. 2003; McLaughlin et al. 2008; Barboza et al. 2017). Symptoms and host(s) The virus induces mosaic, golden yellow, distortion, and stunting symptoms in pepper. The symptom vary based on pepper cultivars and isolates of the virus. Early infected plants produce a few small and distorted fruits causing a serious loss of yield. The Serrano (Ser) strain of PepGMV induced a more chlorotic golden mosaic, whereas the PepGMV-Mo strain induced green-yellow mosaic. The PepGMV-D strain however was very distinct, which induced mild mosaic, distortion, and crumple symptoms only in the inoculated leaves, while newly developed leaves were symptomless (MendezLozano et al. 2003; Brown et al. 2005; Carrillo-Tripp et al. 2007). The natural crop-hosts of PepGMV include cucurbits, pepper, tomato, tomatillo, and tobacco. PepGMV infects tomato in Central America and cucurbits and tomato in Nicaragua (Ala-Poikela et al. 2005). PepGMV infects cucurbits: C. moschata, C. pepo, Sechium edule (chayote) and the Fabaceae plant: Erythrina spp. in Costa Rica (Castro et al. 2013). Transmission The virus is transmitted by the whitefly Bemisia tabaci (Brown and Poulos 1990; Stenger et al. 1990). In common with all other geminiviruses, PepGMV is likely transmitted in a circulative, non-propagative manner. B. tabaci could acquire the virus in 1 h and transmit with a minimum inoculation access period of 48 h. A single whitefly was shown to be able to transmit PepGMV and Pepper huasteco yellow vein virus (PHYVV) simultaneously (Medina-Ramos et al. 2008). PepGMV is mechanically transmissible. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein.

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PepGMV is typical of bipartite begomoviruses native to the New World. The genome consists of two circular, single-stranded DNA components. DNA-A contains 2613 nt (U57457 = NC_004101; AY928516, AY928512, AY928514) and DNA-B of 2595 nt (AF499442 = NC_004096; AY928517, AY928513, AY928515) (Brown and Poulos 1990; Stenger et al. 1990; Torres-Pacheco et al. 1996; Briddon 2001; Brown et al. 2005, 2015; Zerbini et al. 2017). Typical of the majority of bipartite begomoviruses native to the New World the DNA A component of PepGMV encodes six genes, two in the virion-sense and four in the complementary-sense, whereas the DNA B encodes one gene in each orientation. The expression and function of these genes has not been investigated for PepGMV.

Pepper huasteco yellow vein virus

(PHYVV)

Synonyms Pepper hausteco virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution PHYVV infection in plants of Capsicum annuum was reported from Sonora (Mexico), and Texas (USA) (Garzon-Tiznado et al. 1993, Hou et al. 1996; Hernandez-Verdugo et al. 2001; MelendrezBojorquez et al. 2016). Symptoms and host(s) The virus-infected pepper plants show yellow veins initially followed by curling and rugosity of leaves. The symptoms remit later but once the plant is trimmed, the newly developed leaves show severe symptoms (Garzon-Tiznado et al. 1993; Mendez-Lozano et al. 2003). The natural host range of this virus is pepper, tomato, tomatillo, and tobacco. Transmission PHYVV is transmitted by the whitefly Bemisia tabaci. It is likely that, in common with other begomoviruses, transmission is by a circulative, non-propagative pathway. The virus is not transmissible by mechanical inoculation (Medina-Ramos et al. 2008). Virion properties and genome The structure of the virions of PHYVV has not been investigated. In common with all geminiviruses, the virions of PHYVV are likely geminate (twinned quasi-icosahedra). PHYVV is typical of bipartite begomoviruses native to the New World. The genome consists of two circular, single-stranded DNA components. DNA-A contains 2631 nt (X70418 = NC_001359) and DNA-B of 2589 nt (X70419 = NC_001369) (Torres-Pacheco et al. 1993; Briddon 2001; ShimadaBeltran and Rivera-Bustamante 2007; Brown et al. 2015; Zerbini et al. 2017). Typical of the majority of bipartite begomoviruses native to the New World, the DNA A component of PHYVV encodes six genes, one in the virion-sense and four in the complementary-sense, whereas the DNA B encodes one gene in each orientation. The function of these genes has not been investigated for PHYVV.

Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

Pepper leaf curl Bangladesh virus Taxonomic position Genus: Begomovirus

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(PepLCBV)

Family: Geminiviridae

Geographical distribution PepLCBV infection in plants of Capsicum annuum was known to occur in Bangladesh, India, and Pakistan (Tsai et al. unpublished - AF314531). Symptoms and host(s) The virus-infected pepper plants exhibit yellowing and leafcurl symptoms. The host range of the virus has not been investigated. A PCR-based analysis suggested that the virus might also infect tomato. Transmission The transmission of PepLCBV has not been investigated. It is likely that in common with other begomoviruses, PepLCBV is transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Virion properties and genome The structure of the virions of PepLCBV has not been investigated. In common with all geminiviruses, the virions of PepLCBV are likely geminate (twinned quasi-icosahedra). PepLCBV is a typical Old World monopartite begomovirus with a genome that consists of a single circular molecule of single-stranded DNA of ~2753 nt (AF314531 = NC_004192) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes of PepLCBV encode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The function of the genes has not been investigated. The majority of PepLCBV isolates have been shown to be associated with a betasatellite.

Pepper leaf curl Lahore virus Taxonomic position Genus: Begomovirus

(PepLCLaV)

Family: Geminiviridae

Geographical distribution PepLCLaV infection in plants of Capsicum annuum was reported from Pakistan and India (Shafiq et al. 2010; Tahir et al. 2010). Symptoms and host(s) The virus-infected pepper plants exhibit leaf curl symptoms. Transmission The transmission of PepLCLaV has not been investigated. It is likely that, in common with other begomoviruses, PepLCV will be transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Similarly it is likely that the virus will prove not to be either mechanically or seed transmitted.

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Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

Virion properties and genome The structure of the virions of PepLCLaV has not been investigated. In common with all geminiviruses, the virions of PepLCLaV are likely geminate (twinned icosahedra). With only two isolates of PepLCLaV having been characterized, the precise nature of the genome (whether monopartite, monopartite betasatellite-associated or bipartite is unclear). The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2747 nt (AM404179 = NC_016984; JN135234) (Briddon 2001; Shafiq et al. 2010; Tahir et al. 2010; Brown et al. 2015; Zerbini et al. 2017). The genome of PepLCLaV encodes the six genes typically encoded by monopartite begomoviruses and the DNA A component of bipartite begomoviruses. The function of the genes has not been investigated.

Pepper leaf curl virus Taxonomic position Genus: Begomovirus

(PepLCV)

Family: Geminiviridae

Geographical distribution PepLCV infection in plants of Capsicum annuum was known to occur in Malaysia and Thailand (Sinha et al. 2011; Riyaz et al. 2013). Symptoms and host(s) The virus-infected chilli plants exhibit leaf curling symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner (Kumar et al. 2006). The virus is not transmissible by mechanical sap-inoculation. Virion properties and genome The structure of the virions of PepLCV has not been investigated. In common with all geminiviruses, the virions of PepLCV are likely geminate (twinned icosahedra). The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2744 nt (AF134484 = NC_000882). A betasatellite DNA molecule is associated with the virus, and consists of 1361 nt (EF190215) (Briddon 2001; Chattopadhyay et al. 2008; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes/genomic components of PepLCV encode the six genes typically encoded by monopartite begomoviruses and the DNA A component of bipartite begomoviruses. The expression and function of the genes have not been investigated.

Pepper leaf curl Yunnan virus Taxonomic position Genus: Begomovirus

(PepLCYnV)

Family: Geminiviridae

Geographical distribution PepLCYnV infection in plants of Capsicum annuum was reported from Yunnan (China) (Ding et al. unpublished - EU585781).

Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

395

Symptoms and host(s) The virus-infected pepper plants exhibit leaf curling symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm, with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2747 nt (EU585781 = NC_010618) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Pepper leafroll virus Taxonomic position Genus: Begomovirus

(PepLRV)

Family: Geminiviridae

PepLRV infection in plants of Capsicum annuum was reported from Saudi Arabia (Kamran et al. 2018). The virus-infected bell pepper plants show leafroll and interveinal yellowing symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, nonpropagative manner. For more details of PepLRV, refer to Capsicum baccatum cv. Pendulum.

Pepper mild mottle virus Taxonomic position Genus: Tobamovirus

(PMMoV)

Family: Virgaviridae

Geographical distribution PMMoV infection in plants of Capsicum annuum was first described in Italy in 1984 (Wetter et al. 1984). The virus spreads in Argentina, Australia, Egypt, Ethiopia, Senegal, South Africa, Tunisia, Zambia, Taiwan, Korea, Turkey, Denmark, France, Hungary, Iceland, Italy, China, Israel, Japan, the Netherlands, Spain, the UK, Canada, Mexico, Jamaica, Brazil, Argentina, New Zealand, India, Pakistan, and the USA (Wetter et al. 1984; Marte and Wetter 1986; Ndunguru and Kapooria 1996, 1999; Beczner et al. 1997; Adkins et al. 2001; Buzkan et al. 2006; Guldur and Caglar 2006; Wang et al. 2006; CABI/EPPO 2009; Herrera-Vasquez et al. 2009; Caglar et al. 2013; Sevik 2011; Sharma and Patiyal 2011; Ali and Ali 2015; Ahmad et al. 2015; Milosevic et al. 2015; Peng et al. 2015; Rialch et al. 2015; Li et al. 2016; Ruiz et al. 2016; Reddy et al. 2016; Kumar et al. 2017). Symptoms and host(s) The virus-infected pepper plants produce small malformed mottled fruits and sometimes have sunken necrotic spots, mild leaf mottling, and stunting of plants. Symptoms are often more severe in plants infected early, and the severity of symptoms depends on the virulence of the infecting virus strain, the tolerance of the cultivar, and possibly, environmental factors (Wetter et al. 1984). The virus naturally infects bell pepper, chilli, jalapeno pepper, and other Capsicum spp. Several other plant species (Chenopodium spp., Datura stramonium, Nicotiana spp., Ocimum basilicum, Petunia  hybrida, Physalis floridana) are experimentally infected. Tomato is an insusceptible host.

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Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

Transmission The virus is not known to be transmitted by any insect vector. The virus is transmitted by mechanical sap-inoculation, and is transmissible experimentally to at least 24 species of six genera of the Solanaceae and to five species of four genera in four other families, Chenopodiaceae, Cucurbitaceae, Labiatae, and Plantaginaceae. The virus is transmitted by grafting and by contact between plants. PMMoV is seed-borne in Capsicum annuum and C. frutescens up to 100% and the levels of seedborne infection ranged between 0.23% and 100% (Tanzi et al. 1990; Genda et al. 2005; Svoboda et al. 2006). The virus remains infective for prolonged periods both in seed and in soil. The virus has potential to spread through water as genetically diverse PMMoV strains were detected in drinking water sources in Japan (Haramoto et al. 2013). Virion properties and genome The virions are non-enveloped, rigid helical rods with a helical symmetry, about 18 nm in diameter, and 297–312 nm in length. The genome consists of a single molecule of positive-sense ssRNA of 6357 nt (M81413 = NC_003630) (Beczner et al. 1997). The complete genome sequence is available for at least a dozen isolates, which were reported from Brazil, India, Japan, Spain, China, and Korea (Avila-Rincon et al. 1989; Kirita et al. 1997; Kalman et al. 2001; de Oliveira et al. 2010; Rialch et al. 2015). The first complete genome sequence was determined for a resistance-breaking isolate from Spain (PMMoV-S) (Alonso et al. 1991). The genome contains four open reading frames (ORFs) that encode a 126 K protein and a read through 183 K protein, a 28 K movement protein, and a 17.5 K coat protein. PMMoV is one tobamovirus whose ORFs are arranged in non-overlapping manner. There are three phylogenetic clusters of PMMoV isolates. The Spanish isolate S-1a is phylogenetically distinct from the other isolates (de Oliveira et al. 2010). The 30 -terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G50 pppG) (Zaitlin 2011; Adams et al. 2017).

Pepper mottle virus Taxonomic position Genus: Potyvirus

(PepMoV)

Family: Potyviridae

Geographical distribution PepMoV infection in plants of Capsicum annuum was first reported from Florida, USA, by Zitter (1972). The virus has been recorded in Cuba, E1 Salvador, India, Korea, Taiwan, Japan, and the USA (Rodriguez-Alvarado et al. 2002; Ogawa et al. 2003; Warren and Murphy 2003; Choi et al. 2005; Cheng et al. 2011, 2013; Quinones et al. 2011; Kenyon et al. 2014). Symptoms and host(s) The virus causes systemic mild mosaic, vein yellowing, mottling, crinkling, and misshapen leaves and discolored fruits in chilli pepper (Kaur et al. 2014). Transmission The virus is transmitted by aphid vectors, Aphis gossypii, A. craccivora, Myzus persicae in a nonpersistent manner. The virus is transmissible by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants and by seed.

Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

397

Virion properties and genome The virions are non-enveloped, flexuous filaments 737 nm long and 11–13 nm wide. The genome consists of a single-molecule positive-sense ssRNA of 9640 nt (M96425 = NC_001517, AF501591). The genome contains a single long open reading frame starting at nucleotide 168 and potentially encoding a polyprotein of 3068 amino acids (Purcifull et al. 1975; Vance et al. 1992; Warren and Murphy 2003; Revers and Garcia 2015; Wylie et al. 2017).

Pepper necrotic spot virus

(PNSV)

Taxonomic position PNSV is a tentative member of the genus Orthotospovirus and family Tospoviridae Geographical distribution PNSV infection in plants of Capsicum annuum was reported from Peru (Torres et al. 2012). Symptoms and host(s) The virus-infected pepper plants exhibit symptoms of necrotic spots on leaves and stems. Transmission The virus is transmitted by thrips vectors in a persistent, propagative manner. Virion properties and genome The virions are 80–120 nm in diameter. The genome consists of one negative and two ambisense singlestranded RNAs.

Pepper ringspot virus Taxonomic position Genus: Tobravirus

(PepRSV)

Family: Virgaviridae

Geographical distribution PepRSV infection in plants of Capsicum annuum was reported from South and Central American region and Brazil (Robinson 1987). Symptoms and host(s) The virus-infected pepper plants exhibit ringspot symptoms. The virus is known to infect pepper, tomato, and globe artichoke. Transmission The virus is transmitted by a nematode vector, Paratrichodorus minor. The virus is transmissible by mechanical sap-inoculation, grafting but not by contact between plants. Virion properties and genome The virions are rod shaped, non-enveloped, usually straight with a clear modal length of 52 and 197 nm; 22 nm wide. The genome consists of two segments of positive-sense ssRNA (RNA 1 and RNA 2). The RNA 1 and RNA 2 genomes of PepRSV CAM isolate contain 6828 nt (L23972 = NC_003669) and

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Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

1799 nt (X03241 = NC_003670), respectively (Bergh and Siegel 1989; Batista et al. 2014; Rodrigues et al. 2015; Adams et al. 2017).

Pepper severe mosaic virus Taxonomic position Genus: Potyvirus

(PepSMV)

Family: Potyviridae

Geographical distribution PepSMV infection in plants of Capsicum annuum was first reported from Argentina by Feldman and Gracia (1977). The virus spreads in Argentina (Ahn et al. 2006). Symptoms and host(s) The virus-infected pepper plants exhibit symptoms of necrotic spots and streaks in leaves and stems, leaf abscission, then chlorotic leaves may develop. Fruit yield is greatly decreased. Transmission The virus is transmitted by aphid vectors, Aphis craccivora, Macrosiphum euphorbiae and except for type strain, Myzus persicae in a non-persistent manner. The virus is transmissible by mechanical sapinoculation, and also by grafting. Virion properties and genome The virions are flexuous filaments, non-enveloped, with a clear modal length of 761 nm and 13 nm wide. The genome consists of a single molecule of positive-sense ssRNA of 9890 nt (AM181350 = NC_008393) encoding a single polyprotein of 3085 amino acid residues processed into ten functionally distinct proteins (Ahn et al. 2006; Wylie et al. 2017).

Pepper veinal mottle virus Taxonomic position Genus: Potyvirus

(PVMV)

Family: Potyviridae

Geographical distribution PVMV was first reported to be associated with severe diseases in Capsicum annuum and C. frutescens in Ghana (Brunt and Kenten 1971). Subsequently it was reported on peppers in India, Cote d’Ivoire, Nigeria, Mali, Ghana, Kenya, Tunisia, China, Japan, and Africa (de Wijs 1973; Lana et al. 1975a, 1975b; Prasadarao and Yeraguntaiah 1979; Nagaraju and Reddy 1980; Gorsane et al. 1999; Konate and Traore 1999; Gorsane et al. 2001; Moury et al. 2005; Cheng et al. 2009; Tsai et al. 2010; Sharma and Patiyal 2011; Arogundade et al. 2012; Olawale et al. 2015; Matsumoto et al. 2016; Zhang et al. 2016b). Symptoms and host(s) The virus-infected Capsicum spp. plants show vein-clearing followed by systemic interveinal chlorosis, mottling, and distortion of leaves. Fruits of the infected plants are distorted and usually unmarketable. PVMV naturally infects other plant species, e.g., tomato, tobacco, eggplant, petunia, Solanum nigrum, Datura metel, D. stramonium, Physalis angulata, P. micrantha, and Telfairia occidentalis (Brunt et al. 1978; Arogundade et al. 2012).

Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

399

Transmission The virus is transmitted by aphid vectors such as, Aphis gossypii, A. spiraecola, Myzus persicae, and Toxoptera citricidus in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation to a number of cultivated and weed hosts. The virus is transmissible by grafting. The virus is not transmissible by contact between plants. The virus is not seed-borne in pepper. Virion properties and genome The virions are non-enveloped, flexuous filaments 770 nm long and 12 nm wide. The genome consists of a single molecule of positive-sense ssRNA of 9792 nt with a poly-A tail (DQ645484 = NC_011918) (Moury et al. 2005; Ha et al. 2008; Revers and Garcia 2015; Wylie et al. 2017).

Pepper vein yellows virus 1

(PeVYV-1)

Synonyms Pepper vein yellows virus (PeVYV); Pepper yellows virus Taxonomic position Genus: Polerovirus

Family: Luteoviridae

Geographical distribution PeVYV-1 infection in plants of Capsicum annuum and Capsicum frutescens was reported from the Pakistan, the USA, Sudan, Japan, China, Spain, Italy, Ivory Coast, Benin, and Israel (Yonaha et al. 1995; Villanueva et al. 2013; Alfaro-Fernandez et al. 2014; Alabi et al. 2015; Bolou Bi et al. 2015; Tan et al. 2015; Zhang et al. 2015a; Liu et al. 2016; Tomassoli et al. 2016; Afouda et al. 2017; Lotos et al. 2017; Murakami and Kawano 2017; Ahmad et al. 2018). Symptoms and host(s) The virus-infected pepper plants exhibit symptoms of interveinal yellowing, upward leaf curling, internode shortening, and abnormal ripening of fruits. Transmission The virus is transmitted by aphid vectors, Myzus persicae and Aphis gossypii in a circulative and nonpropagative manner. Virion properties and genome The virions are 25–30 nm in diameter and hexagonal in outline and have no envelope. The genome consists of a single molecule of infectious, linear, positive-sense ssRNA of 6244 nt (AB594828). The 30 terminus has neither a poly (A) tract nor a tRNA-like structure. The 50 terminus has a genome-linked protein (VPg) (of about 17 kDa). The genome contains six open reading frames (ORFs) (Yonaha et al. 1995; Murakami et al. 2011; Liu et al. 2016).

Pepper vein yellows virus 2 Synonyms Pepper yellow leaf curl virus

(PeVYV-2)

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Taxonomic position Genus: Polerovirus

Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

Family: Luteoviridae

Geographical distribution PeVYV-2 infection in plants of Capsicum annuum was reported from Israel (Dombrovsky et al. 2013). Symptoms and host(s) The virus-infected pepper plants exhibit vein yellowing and leaf roll symptoms. Transmission The virus is transmitted by aphid vectors in a circulative and nonpropagative manner. Virion properties and genome The virions are 25–30 nm in diameter, hexagonal in outline, and have no envelope. The genome consists of a single molecule of infectious, linear, positive-sense ssRNA of 6028 nt (HM439608) and encoding six open reading frames (ORFs) (Dombrovsky et al. 2013).

Pepper vein yellows virus 3 Taxonomic position Genus: Polerovirus

(PeVYV-3)

Family: Luteoviridae

Geographical distribution PeVYV-3 infection in plants of Capsicum annuum was reported from China (Liu et al. 2016). Symptoms and host(s) The virus-infected pepper plants exhibit vein yellowing and leaf roll symptoms. Transmission The virus is transmitted by aphid vectors in a circulative and nonpropagative manner. Virion properties and genome The virions are 25–30 nm in diameter, hexagonal in outline, and have no envelope. The genome consists of a single molecule of infectious, linear, positive-sense ssRNA of 6244 nt (KP326573), and contains six open reading frames (ORFs) (Liu et al. 2016).

Pepper vein yellows virus 4 Taxonomic position Genus: Polerovirus

(PeVYV-4)

Family: Luteoviridae

Geographical distribution PeVYV-4 infection in plants of Capsicum annuum was reported from Australia (Maina et al. 2016). Symptoms and host(s) The virus-infected pepper plants exhibit vein yellowing and leaf roll symptoms.

Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

401

Transmission The virus is transmitted by aphid vectors in a circulative and nonpropagative manner. Virion properties and genome The virions are 25–30 nm in diameter, hexagonal in outline, and have no envelope. The genome consists of a single molecule of infectious, linear, positive-sense ssRNA of 6244 nt (KU999109), with the six open reading frames (ORFs), ORF0 and ORF2 overlap with ORF1, and ORF4 is included within ORF3 to ORF5 (Maina et al. 2016).

Pepper vein yellows virus 5 Taxonomic position Genus: Polerovirus

(PeVYV-5)

Family: Luteoviridae

Geographical distribution PeVYV-5 infection in plants of Capsicum annuum cv. California Wonder was reported from Spain (Fiallo-Olivé et al. 2018). Symptoms and host(s) The virus-infected pepper plants exhibit vein yellowing and leaf roll symptoms. Transmission The virus is transmitted by aphid vector Aphis gossypii in a circulative and nonpropagative manner. Virion properties and genome The virions are 25–30 nm in diameter, hexagonal in outline, and have no envelope. The genome consists of a single molecule of infectious, linear, positive-sense ssRNA of 6125 nt (KY523072) and contains seven open reading frames (ORF0 to ORF5, and ORF3a) (Fiallo-Olivé et al. 2018).

Pepper vein yellows virus 6 Taxonomic position Genus: Polerovirus

(PeVYV-6)

Family: Luteoviridae

Geographical distribution PeVYV-6 infection in plants of Capsicum annuum was reported from Greece (Lotos et al. 2017). Symptoms and host(s) The virus-infected pepper plants exhibit symptoms of interveinal yellowing, severe upward leafroll, and stunted growth. Transmission The virus is transmitted by aphid vectors in a circulative and nonpropagative manner.

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Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

Virion properties and genome The virions are 25–30 nm in diameter, hexagonal in outline, and have no envelope. The genome consists of a single molecule of infectious, linear, positive-sense ssRNA of 6096 nt (LT559483) and containing seven open reading frames (ORF0 to ORF5, and ORF3a) (Lotos et al. 2017).

Pepper yellow leaf curl Indonesia virus Taxonomic position Genus: Begomovirus

(PepYLCIV)

Family: Geminiviridae

Geographical distribution PepYLCIV infection in plants of Capsicum annuum was reported from Indonesia (Tsai et al. 2006). Symptoms and host(s) The virus-infected pepper plants show yellow leaf curl symptoms (Tsai et al. 2006). The virus has been isolated from chilli pepper (Capsicum annuum), tomato (Solanum lycopersicum), and Ageratum conyzoides. Transmission The transmission of PepYLCIV by Bemisia tabaci, the whitefly vector of begomoviruses, has been demonstrated (Sulandari et al. 2007). The mechanism of transmission has not been investigated, but will likely be in a circulative, non-propagative manner. Experimentally, the virus has been shown to infect Nicotiana benthamiana, Nicotiana glutinosa, Nicotiana tabacum, Vigna unguiculata, Vigna radiata, Glycine max, and Physalis floridana by Bemisia tabaci transmission (Sulandari et al. 2007). Virion properties and genome The structure of PepYLCIV virions has not been investigated. In common with all geminiviruses, the virions of PepYLCIV are likely geminate (twinned icosahedra). The genome of PepYLCIV is bipartite consisting of two circular, single-stranded DNA components. DNA-A contains 2750 nt (AB267834 = NC_008283; AB267836, AB267838, DQ083764, DQ083765) and DNA-B of 2726 nt (AB267835 = NC_008284; AB267837, AB267839) (Briddon 2001; Jamsari and Pedri 2013; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the Old World, the DNA A component of PepYLCIV encodes six genes, two in the virion-sense and four in the complementary-sense.

Pepper yellow leaf curl Indonesia virus 2 Taxonomic position Genus: Begomovirus

(PepYLCIV2)

Family: Geminiviridae

Geographical distribution PepYLCIV2 infection in plants of Capsicum annuum was reported from Indonesia (Jamsari et al. unpublished – KT809345).

Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

403

Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, nonpropagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2748 nt (KT809345) (Brown et al. 2015; Zerbini et al. 2017).

Pepper yellow leaf curl Thailand virus Taxonomic position Genus: Begomovirus

(PepYLCTHV)

Family: Geminiviridae

Geographical distribution PepYLCTHV infection in plants of Capsicum annuum was reported from Thailand (Yule et al. unpublished – KX943290, KX943291; Chiemsombat et al. 2018) Symptoms and host(s) The virus-infected pepper plants show striking yellow mosaic and leaf curl, or yellow veins and leaf curl. Young leaves were chlorotic and shoots were severely distorted Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is bipartite consisting of two circular, single stranded DNA components. DNA-A consists of 2742 nt (KX943290) and DNA-B of 2732 nt (KX943291) (Briddon 2001; Brown et al. 2015; Chiemsombat et al. 2018; Zerbini et al. 2017).

Pepper yellow leaf curl virus Taxonomic position Genus: Begomovirus

(PepYLCV)

Family: Geminiviridae

Geographical distribution PepYLCV infection in plants of Capsicum annuum was reported from China, Israel, and Indonesia (De Barro et al. 2008; Dombrovsky et al. 2010). Symptoms and host(s) The virus-infected pepper plants exhibit symptoms of shortening of stem internodes, interveinal yellowing, and upward rolling of the leaf blade, accompanied by fruit discoloration and size reduction (Dombrovsky et al. 2010).

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Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner (De Barro et al. 2008). Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2748 nt (KC149938 = NC_020236) (Briddon 2001; Dombrovsky et al. 2013; Brown et al. 2015; Zerbini et al. 2017).

Pepper yellow mosaic virus Taxonomic position Genus: Potyvirus

(PepYMV)

Family: Potyviridae

Geographical distribution PepYMV infection in plants of Capsicum annuum was widely distributed in Brazil (Inoue-Nagata et al. 2002, 2003; Cunha et al. 2004; Truta et al. 2004; Moura et al. 2011; Lucinda et al. 2012). Symptoms and host(s) The virus-infected pepper plants exhibit mosaic, yellow mosaic, vein-banding, leaf distortion, blistering and stunting. The virus infects tomato, sweet pepper, and chili pepper in Brazil. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically saptransmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments 770 nm long and 13 nm wide. The genome of contains a single molecule of positive-sense ssRNA of 9745 nt, excluding the 30 poly(A) tail (AB541985 = NC_014327) (Cunha et al. 2004; Lucinda et al. 2012; Revers and Garcia 2015; Wylie et al. 2017).

Pepper yellow vein Mali virus Taxonomic position Genus: Begomovirus

(PepYVMLV)

Family: Geminiviridae

Geographical distribution PepYVMLV in plants of Capsicum annuum was reported from Mali, Burkina Faso, Cote d’Ivoire, and China (Tiendrebeogo et al. 2008, 2011; Seka et al. 2017). Symptoms and host(s) The virus-infected pepper plants exhibit severe foliar yellowing, leaf curling, and leaf distortion symptoms.

Capsicum annuum and Capsicum frutescens (Bell pepper, Chilli, Pepper, Sweet pepper)

405

The virus has been detected based on complete genome sequences from different weed and crop species in Fujian, China, that include Eclipta prostrata (AM691547), Phyllanthus urinaria (AM691550), Bougainvillea (AM691549), and Capsicum (AM691548). Transmission The virus was shown to be transmitted by the whitefly Bemisia tabaci (Tiendrebeogo et al. 2008, 2011). The mechanism of transmission was not investigated. It is likely that in common with other begomoviruses, PepYVMLV is transmitted by B. tabaci in a circulative, non-propagative manner. Experimentally, the virus has been introduced into Nicotiana benthamiana. The virus was not seed transmissible. Virion properties and genome The structure of the virions of PepYVMLV has not been investigated. In common with all geminiviruses, the virions of PepYVMLV are likely geminate (twinned quasi-icosahedra). PepYVMLV is a typical Old World monopartite begomovirus with a genome consisting of a single circular molecule of single-stranded DNA of ~2786 nt (AY502935 = NC_005347; FN555172; AM691547, AM691548, AM691549, AM691550, AM691555, FM876847, FM876848, FM876849, FM876850, FM876851, FM876852, FN555171, FN555173, FN555174) (Briddon 2001; Tiendrebeogo et al. 2011; Brown et al. 2015). The characterized genomes of PepYVMLV encode the six genes typically encoded by monopartite begomoviruses originating from the Old World, two in the virion-sense and four in the complementary-sense. The expression and function of the genes have not been investigated.

Potato spindle tuber viroid Taxonomic position Genus: Pospiviroid

(PSTVd)

Family: Pospiviroidae

PSTVd infection in plants of Capsicum annuum was reported from New Zealand, the Netherlands, and Australia (Lebas et al. 2005; Mackie et al. 2016). The viroid-infected pepper plants exhibit interveinal chlorosis, epinasty, and brittleness symptoms. The viroid is mechanically sap-transmissible and also spreads easily through use of contaminated tools. The viroid is seed-transmitted through the seeds of Capsicum annuum var grossum to the extent of 0.3% (Matsushita and Tsuda 2016). For more details of PSTVd, refer to Solanum tuberosum.

Potato virus Y

(PVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

PVY has been reported to infect Capsicum annuum in India, Sudan, South Africa, Spain, Tehran Province of Iran, the United States, Zambia, Zimbabwe, Serbia, the Czech Republic and most other peppergrowing regions (George et al. 1993; Luis-Arteaga et al. 1997; Ndunguru and Kapooria 1999; LuisArteaga and Ponz, 2003; Fanigliulo et al. 2005; Jagadeeshwar et al. 2005; Mostafae et al. 2006, 2012; Petrovic et al. 2010; Sharma and Patiyal 2011; Svoboda and Svobodova-Leisova 2012; Moodley et al. 2014; Olawale et al. 2015; Karavina et al. 2016). The virus-infected pepper plants exhibit symptoms of

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mild to severe leaf mosaic, narrow bands of dark green tissue along the leaf veins. Some PVY strains on pepper produce veinal necrosis symptoms (Mills and Abdul-Magid 1987; d’Aguino et al. 1995). PVY pepper isolates have been classified into three pathotypes: PVY-0, PVY-1, and PVY-1-2, based on their ability to overcome certain resistance genes in peppers. Pathotype PVY-0 occurs worldwide, while the other two pathotypes occur predominantly in tropical and subtropical countries (GebreSelassie et al. 1983). The pepper isolates of PVY were different from potato isolates of PVY in host range, aphid transmission efficiencies, serology, and sequence of coat protein (Romero et al. 2001; Janzac et al. 2010). The virus is transmitted by aphid vectors, viz., Myzus persicae, Macrosiphum euphorbiae, and other aphid species spreads the virus from plant to plant during very short feeding probes. The virus is mechanically sap-transmissible to number of hosts. Otulak et al. (2017) have demonstrated PVYNTN transmission through C. annuum seeds, and they have observed virus particles inside seeds on embryo surfaces, while particles and Potyvirus inclusion bodies in endothelium layers. For more details of PVY, refer to Solanum tuberosum.

Rehmannia mosaic virus Taxonomic position Genus: Tobamovirus

(RheMV)

Family: Virgaviridae

RheMV infection in plants of Capsicum annuum was reported from Japan (Kubota et al. 2012). The virus-infected pepper plants exhibit symptoms of severe necrotic mosaic. As the symptoms progressed, the infected plants developed prominent leaf necrosis and severe leaf fall, stem necrosis, and fruit distortion. The virus is mechanically sap-transmissible and also through contact between plants and through seed (Kubota et al. 2012). For more details of RheMV, refer to Rehmannia glutinosa.

Sri Lankan cassava mosaic virus Taxonomic position Genus: Begomovirus

(SLCMV)

Family: Geminiviridae

SLCMV infection in plants of Capsicum annuum was reported from India (Khan et al. 2011). The virusinfected pepper plants exhibit mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of SLCMV, refer to Manihot esculenta.

Tobacco curly shoot virus Taxonomic position Genus: Begomovirus

(TbCSV)

Family: Geminiviridae

TbCSV infection in plants of Capsicum frutescens was reported from China (Qing et al. 2010). The virus-infected bell pepper plants exhibit stunting, leaf yellowing, and mild curling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. For more details of TbCSV, refer to Nicotiana tabacum.

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Tobacco etch virus Taxonomic position Genus: Potyvirus

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(TEV)

Family: Potyviridae

TEV infection in plants of Capsicum annuum was reported from India, the Americas, Caribbean, Jamaica, Sudan, Cuba, and Turkey (Bidari and Reddy 1986, 1990; Mills and Abdul-Magid 1987; Depestre et al. 1993; Chu et al. 1997; Pernezny et al. 2003; Buzkan et al. 2006; Olawale et al. 2015). The virus-infected pepper plants exhibit symptoms of mottling, distortion, downward curling, vein-clearing, necrotic lines, and mosaic pattern symptoms on the leaves. The virus is transmitted by aphid vectors such as Myzus persicae, and Aphis gossypii in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TEV, refer to Nicotiana tabacum.

Tobacco mild green mosaic virus Taxonomic position Genus: Tobamovirus

(TMGMV)

Family: Virgaviridae

TMGMV infection in plants of Capsicum annuum occurs in worldwide, including Italy, Korea, Taiwan, Tunisia, Panama, Asia and Central/South America, Hungary, and Japan (Takeuchi et al. 1998; Choi et al. 2002, 2005; Li and Chang 2005; Herrera-Vasquez et al. 2009; Font et al. 2009; Chen et al. 2013; Nemes et al. 2015). Symptoms on Capsicum annuum vary with the pepper genotype and environmental conditions. The virus-infected pepper plants exhibit mild green and chlorotic mosaic symptoms or even chlorotic ringspots and severe necrotic mosaic by which plants are often killed. No insect vector is reported for this virus. The virus is transmissible by mechanical sap-inoculation, and has a wide host range. The virus is transmitted by grafting but not through seed. For more details of TMGMV, refer to Nicotiana tabacum.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV infection in plants of Capsicum annuum was reported from India, southern Europe, Nigeria, Zambia, the USA, and China (Nagai et al. 1981; Sulaiman and Gim 1981; Igwegbe 1983; Avgelis 1986; Sakata et al. 1997; Chitra et al. 1999; Ndunguru and Kapooria 1999; Murphy et al. 2003; Rao and Lan 2003; Jagadeeshwar et al. 2005; Sharma and Patiyal 2011; Moury and Verdin 2012; Pazarlar et al. 2013; Kumar et al. 2017). The virus-infected pepper plants show various degrees of mosaic, and/or necrosis on leaves, stems, and fruits and stunting in pepper. The virus is more common in pepper under protected cultivation. The effects of TMVon the greenhouse pepper cvs “Ergenekon F1,” “Kumsal F1,” and 497 F1 in Turkey include reduction of vegetative growth of chlorophyll a and b, plant fresh, and dry production, but increase of proline content in leaves, although the level varied from variety to variety (Pazarlar et al. 2013). The virus may be introduced into fields in transplants, in crop debris, and on hands and clothing of workers, as well as on contaminated tools and machinery (Kim et al. 1989). Workers can get TMVladen sap from infected plants on their hands and readily transmit the virus from plant to plant down the row. Some tobacco products used by workers would be a very important source of TMV. The virus is

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highly contagious and also transmitted through seeds of chilli to the extent of 13.5–29.6% (Tosic et al. 1980; Demski 1981; Chitra et al. 1998, 2002; Kumar and Prakash 2016), but no transmission through pollen. No insect vector is reported. For more details of TMV, refer to Nicotiana tabacum.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in plants of Capsicum annuum was reported from Japan, Brazil, China, the USA, and Italy (Semancik 1966; Tian and Zhang 1995; Tomitaka et al. 2011). The virus-infected pepper plants exhibit severe chlorosis and necrosis symptoms. The virus is transmitted by nematode vectors, and Trichodorus and/or Paratrichodorus which are shown to spread TRV to pepper transplants through contaminated soil. The virus is transmissible by mechanical sap-inoculation to a large number of host plants. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Capsicum annuum was reported from India, Australia, and Argentina (Gracia and Feldman 1974; Jain et al. 2005; Sharman et al. 2015). The virus-infected pepper plants exhibit symptoms of systemic necrosis, especially of the upper leaves, dark streaks on stems and petioles, and leaf drop. The virus is transmitted by the thrips vectors, presents in/on pollen, and enters in to the host through the injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible and is not transmissible by contact between plants (Sdoodee and Teakle 1987). For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tobacco yellow crinkle virus Taxonomic position Genus: Begomovirus

(TbYCV)

Family: Geminiviridae

TbYCV infection in plants of Capsicum annuum was reported from Cuba (Fiallo-Olive et al. 2009). The virus-infected pepper plants exhibit foliar yellow mottle symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of TbYCV, refer to Nicotiana tabacum.

Tomato apical stunt viroid Taxonomic position Genus: Pospiviroid

(TASVd)

Family: Pospiviroidae

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TASVd infection in plants of Capsicum annuum was reported from Taiwan (Verhoeven et al. 2017). The viroid is mechanically sap-transmissible and also through seed. For more details of TASVd, refer to Solanum lycopersicum.

Tomato aspermy virus

(TAV)

Synonyms Chrysanthemum aspermy virus; Chrysanthemum mild mottle virus Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

TAV infection in plants of Capsicum annuum was reported from Japan (Natsuaki et al. 1994). The virusinfected pepper plants exhibit necrotic mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TAV, refer to Solanum lycopersicum.

Tomato bushy stunt virus Taxonomic position Genus: Tombusvirus

(TBSV)

Family: Tombusviridae

TBSV infection in plants of Capsicum annuum spreads in Argentina, Morocco, Tunisia, the UK, and the USA (Fischer and Lockhart 1977; Borges et al. 1979; Cherif and Spire 1983). The virus-infected pepper plants exhibit severe symptoms such as stunting, leaf deformation, strong mottle, flower abscission, and lethal systemic necrosis. The virus is transmissible by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants. There is no known vector for this virus. For more details of TBSV, refer to Solanum lycopersicum.

Tomato chino La Paz virus Taxonomic position Genus: Begomovirus

(ToChLPV)

Family: Geminiviridae

ToChLPV infection in plants of Capsicum annuum var. ancho poblano was reported from the State of Baja California Sur, Mexico (Cardenas-Conejo et al. 2010). The virus-infected pepper plants exhibit yellowing and stunting of plants and foliage discoloration and flower abortion symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ToChLPV, refer to Solanum lycopersicum.

Tomato chlorosis virus Taxonomic position Genus: Crinivirus

(ToCV)

Family: Closteroviridae

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ToCV infection in plants of Capsicum annuum was reported from Spain, Costa Rica, Tunisia, Uruguay, and Brazil (Lozano et al. 2004; Barbosa et al. 2010; Vargas et al. 2011; Fortes et al. 2012). The virusinfected pepper plants exhibit symptoms of interveinal yellowing, leaf curling, and stunting. The virus is transmitted by whitefly vectors, Bemisia tabaci biotype Q and Trialeurodes vaporariorum in a semipersistent manner (Vargas-Asencio et al. 2013). For more details of ToCV, refer to Solanum lycopersicum.

Tomato chlorotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(TCSV)

Family: Tospoviridae

TCSV infection in plants of Capsicum annuum was reported from the United States, Brazil, Puerto Rico, Cuba, and Argentina (Boiteux et al. 1993; Gracia et al. 1999; Dal Bo et al. 2001; Almeida et al. 2014; Webster et al. 2013, 2015; Gonzalez-Alvarez et al. 2017). The virus-infected pepper plants exhibit symptoms of necrotic spots or ringspots and severe leaf deformation, if infected at a young stage, they fail to grow and are severely stunted. The fruits from plants that are infected later are deformed and discolored and show ringspot symptoms. The virus is transmitted by a thrips vector, Frankliniella schultzei, in a persistent-propagative manner, and is also by mechanical sap-inoculation. For more details of TCSV, refer to Solanum lycopersicum.

Tomato dwarf leaf virus Taxonomic position Genus: Begomovirus

(ToDfLV)

Family: Geminiviridae

ToDfLV has been reported to infect plants of Capsicum annuum in Jamaica (Roye et al. 1999). The virus causes severe dwarf and leaf curling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ToDfLV, refer to Solanum lycopersicum.

Tomato leaf curl Cebu virus Taxonomic position Genus: Begomovirus

(ToLCCeV)

Family: Geminiviridae

ToLCCeV infection in plants of Capsicum annuum was reported from the Philippines (Tsai et al. 2011). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ToLCPV, refer to Capsicum annuum.

Tomato leaf curl Joydebpur virus Taxonomic position Genus: Begomovirus

(ToLCJoV)

Family: Geminiviridae

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ToLCJoV infection in plants of Capsicum annuum was reported from India (Shih et al. 2007). The virus-infected pepper plants exhibit mild yellowing, severe leaf curling, leaf distortion, stunting, and blistering symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner, and the virus is also graft-transmissible. Tomato leaf curl Joydebpur betasatellite DNA molecule is associated with ToLCJoV, and consists of 1362 nt (HM007103) is associated with ToLCJoV (Zhou 2013). For more details of ToLCJoV, refer to Solanum lycopersicum.

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Capsicum annuum was reported from India and Pakistan (Khan et al. 2006, 2014; Khan and Raj 2008; Hussain et al. 2004). The virus-affected pepper plants often show symptoms consisting of leaf curling, shortening of internodes and petioles, crowding of leaves, and stunting of the whole plant. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is graft-transmissible and not through seed. For more details of ToLCNDV, refer to Solanum lycopersicum.

Tomato leaf curl Philippines virus Taxonomic position Genus: Begomovirus

(ToLCPV)

Family: Geminiviridae

ToLCPV infection in plants of Capsicum annuum was reported from the Philippines (Tsai et al. 2011). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ToLCPV, refer to Solanum lycopersicum.

Tomato leaf curl Sinaloa virus Taxonomic position Genus: Begomovirus

(ToLCSiV)

Family: Geminiviridae

ToLCSiV infection in plants of Capsicum annuum was reported from Mexico (Brown et al. 1993). The virus-infected pepper plants exhibit splotchy green mottle symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is graft-transmissible and not through seed. For more details of ToLCSiV, refer to Solanum lycopersicum.

Tomato leaf curl Sulawesi virus Taxonomic position Genus: Begomovirus

(ToLCSuV)

Family: Geminiviridae

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ToLCSuV infection in plants of Capsicum annuum was reported from Sulawesi (Indonesia) (Tsai et al. 2009). The virus-infected pepper plants exhibit leaf curling, blistering, yellowing, and stunting symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative nonpropagative manner. For more details of ToLCSuV, refer to Solanum lycopersicum.

Tomato marchitez virus Taxonomic position Genus: Torradovirus

(ToMarV)

Family: Secoviridae

ToMarV infection in plants of Capsicum annuum was reported from Mexico (Camacho-Beltran et al. 2015). The virus-infected pepper plants exhibit symptoms of yellow mosaic, upward leaf curling, crinkling, and stunting. The virus is transmitted by whitefly vectors in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of ToMarV, refer to Solanum lycopersicum.

Tomato mosaic virus Taxonomic position Genus: Tobamovirus

(ToMV)

Family: Virgaviridae

ToMV infection in plants of Capsicum annuum was reported from southern Europe and Syria (Ahoonmanesh et al. 1992; Chitra et al. 1999; Ismaeil et al. 2012; Moury and Verdin 2012; Olawale et al. 2015; Kumar et al. 2017). The virus-infected pepper plants exhibit symptoms of leaf distortion with chlorotic mottle or mosaic and small, misshapen, and discolored fruits. Often there are necrotic patches on fruits and leaves. No vector is involved in the spread of this virus. The virus is mechanically sap-transmissible and also transmissible through contact between plants. The virus is seed-transmitted through the seeds of Capsicum annuum to the extent of 11.5% (Kumar and Prakash 2016). For more details of ToMV, refer to Solanum lycopersicum.

Tomato mottle mosaic virus Taxonomic position Genus: Tobamovirus

(ToMMV)

Family: Virgaviridae

ToMMV infection in plants of Capsicum annuum var. grossum was reported from China (Li et al. 2014). The virus-infected pepper plants exhibit symptoms of mottling, stunting, and necrosis symptoms. No vector is known for this virus. The virus is transmissible by mechanical sap-inoculation, and also through contact between plants. For more details of ToMMV, refer to Solanum lycopersicum.

Tomato necrotic ringspot virus

(TNRV)

Taxonomic position TNRV is a tentative member of the genus Orthotospovirus and family Tospoviridae

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TNRV infection in plants of Capsicum frutescens was reported from Thailand (Chiemsombat et al. 2010; Hassani-Mehraban et al. 2011). The virus-infected bell pepper plants exhibit severe necrotic mottling and/or necrotic spots and rings on the leaves and fruits of the respective plants. The virus could be transmitted by Thrips palmi and Ceratothripoides claratris, with T. palmi being the most efficient vector for this virus. The virus is mechanically sap-transmissible. For more details of TNRV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV was identified to occur in tabasco (Capsicum frutescens) peppers in several districts of Tehran, northern Iran, and the USA (Sokhansanj et al. 2012). The infected bell pepper plants exhibit mosaic, chlorosis, and malformation symptoms on leaves as well as on fruits. The virus is transmitted by a nematode vector, Xiphinema americanum (sensu lato), in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and also by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Tomato severe rugose virus Taxonomic position Genus: Begomovirus

(ToSRV)

Family: Geminiviridae

ToSRV infection in plants of Capsicum annuum was reported from Brazil (Bezerra-Agasie et al. 2006; Nozaki et al. 2006; Nozaki et al. 2010; Rocha et al. 2010). The virus-infected pepper plants exhibit yellow mosaic and leaf distortion symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner to tomato and pepper. For more details of ToSRV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infects Capsicum annuum plants in different parts of the world (Reddick et al. 1987; de Avila et al. 1991; Gaborjanyi et al. 1995; Skoric et al. 1997; Yurtmen et al. 1999; Gera et al. 2000; Momol et al. 2000; Kim et al. 2004; Crescenzi et al. 2013; Petrovic et al. 2010; Zheng et al. 2010; Sharma and Patiyal 2011; Zindovic et al. 2011; Deligoz et al. 2014; Martinez et al. 2014; Perez-Colmenares et al. 2015; Karavina et al. 2016; Ashfaq and Ahmad 2017; Karavina and Gubba 2017; Sivaprasad et al. 2017; Delic et al. 2018). The virus-infected pepper plants exhibit initial symptoms on leaves of small necrotic lesions that coalesce with time, often accompanied by ringspots. Early infection may lead to severe stunting of the plant and in combination with the droopy leaves, give the impression that they are wilted. Symptoms on the green fruits are concentric rings of yellow or brown alternating with the background green color, or striking brown rings occurring on red-ripe fruit.

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This virus spreads primarily by the insect thrips, Frankliniella occidentalis, and also by F. bispinosa in a persistent-propagative manner (Avila et al. 2006). The virus is mechanically sap-transmissible to number of hosts. The virus is not seed-borne, does not spread by contact and does not survive in soil or debris. For more details of TSWV, refer to Solanum lycopersicum.

Tomato torrado virus Taxonomic position Genus: Torradovirus

(ToTV)

Family: Secoviridae

ToTV infection in plants of Capsicum annuum was reported from Europe, Central America, and Australia (Amari et al. 2008). The virus-infected pepper plants show severe symptoms of conspicuous mosaic and/or stunting but no necrosis. The virus is transmitted by the whitefly vectors, Bemisia tabaci, and also by Trialeurodes vaporariorum in a semi-persistent manner, and also by mechanical sapinoculation. For more details of ToTV, refer to Solanum lycopersicum.

Tomato yellow leaf curl Mali virus Taxonomic position Genus: Begomovirus

(TYLCMLV)

Family: Geminiviridae

TYLCMLV infection in plants of Capsicum annuum was reported from Cameroon (Lett et al. 2009). The virus-infected pepper plants exhibit symptoms of stunting or dwarfing, with leaflets rolled upwards and inwards; young leaves are slightly chlorotic. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not transmitted by contact between plants. For more details of TYLCMLV, refer to Solanum lycopersicum.

Tomato yellow leaf curl Thailand virus Taxonomic position Genus: Begomovirus

(TYLCTHV)

Family: Geminiviridae

TYLCTHV infection in plants of Capsicum annuum was reported from Taiwan (Shih et al. 2010). The virus-infected pepper plants exhibit leaf curling, blistering, distortion, mild vein-yellowing, and stunting symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative nonpropagative manner. The virus is transmissible by grafting but not transmitted by contact between plants. For more details of TYLCTHV, refer to Solanum lycopersicum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV has been found to infect plants of Capsicum annuum in China, Mexico, Oman, Spain, the Dominican Republic, Egypt, Morocco, Florida, Cuba, Guatemala, Jamaica, Korea, India, and Tanzania

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(Shalaby et al. 1997; Reina et al. 1999; Quinones et al. 2002; Morilla et al. 2005; Polston et al. 2006; Chouchane et al. 2007; Salati et al. 2010; Sharma and Patiyal 2011; Kil et al. 2014). The virus-infected pepper plants are often symptomless (Morilla et al. 2005). Although many pepper cultivars appear susceptible to TYLCV, large differences in infection rates have been observed, both under field conditions and by inoculation under controlled conditions. Infected pepper plants (Capsicum annuum) show mild interveinal yellowing and curling of leaves (Quinones et al. 2002). The virus is transmitted by the Q biotype of Bemisia tabaci in a circulative non-propagative manner. The virus is transmissible by grafting but not by contact between plants (Morilla et al. 2005) and is seed-transmitted in C. annuum (Kil et al. 2018). For more details of TYLCV, refer to Solanum lycopersicum.

Tomato yellow ring virus

(TYRV)

Synonyms Tomato yellow fruit ring virus (TYFRV) Taxonomic position TYRV is a tentative member of the genus Orthotospovirus and family Tospoviridae TYFRV infection in plants of Capsicum annuum was reported from Iran (Golnaraghi et al. 2013). The virus-infected pepper plants exhibit yellow mosaic, chlorosis, and necrosis symptoms. The virus is transmitted by the thrips vector, Thrips tabaci, in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of TYRV, refer to Solanum lycopersicum.

Tomato yellow vein streak virus Taxonomic position Genus: Begomovirus

(ToYVSV)

Family: Geminiviridae

ToYVSV infection in plants of Capsicum annuum was reported from Brazil (Nozaki et al. 2010; InoueNagata et al. 2016). The virus is easily transmitted by the whitefly Bemisia tabaci biotype B in a circulative non-propagative manner. The virus is mechanically sap-transmissible. For more details of ToYVSV, refer to Solanum lycopersicum.

Tomato zonate spot virus

(TZSV)

Taxonomic position TZSV is a tentative member of the genus Orthotospovirus and family Tospoviridae TZSV infection in plants of Capsicum annuum was reported from China (Dong et al. 2008). The virusinfected pepper plants exhibit initial symptoms including internal necrotic lesions of the veins, petioles, and stems. As symptoms became fully developed, the internal necrotic lesions in the stem enlarged, which resulted in the death of infected plants. Even chlorotic or necrotic flecking on leaves and chlorotic ringspots on fruit may be noticed. The virus is transmitted by thrips vectors in a persistent propagative manner. For more details of TZSV, refer to Solanum lycopersicum.

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Watermelon bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(WBNV)

Family: Tospoviridae

WBNV infection in plants of Capsicum annuum was reported from India (Bhat et al. 2014). The virus is successfully transmitted by the thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of WBNV, refer to Citrullus lanatus.

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV infection in plants of Capsicum annuum was reported from Iran (Hajizadeh and Mohammadi 2016). The virus-infected pepper plants exhibit mosaic, leaf distortion, mottling, stunting, and yellowing symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of WMV, refer to Citrullus lanatus.

White clover mosaic virus Taxonomic position Genus: Potexvirus

(WClMV)

Family: Alphaflexiviridae

WClMV infection in plants of Capsicum annuum was reported from Saudi Arabia (Al-Shahwan et al. 2017). The virus is sap-transmissible and also through contact between plants. For more details of WClMV, refer to Trifolium spp.

Youcai mosaic virus Taxonomic position Genus: Tobamovirus

(YoMV)

Family: Virgaviridae

YoMV infection in plants of Capsicum annuum was reported from China (Zhang et al. 2016a). The virus-infected pepper plants exhibit chlorotic and mild upward leaf curling symptoms. There is no known vector for this virus. The virus is mechanically sap-transmissible and also transmissible through contact between plants. For more details of YoMV, refer to Brassica napus.

References Abdalla OA, Ali A (2012) First report of Alfalfa mosaic virus associated with severe mosaic and mottling of pepper (Capsicum annuum) and white clover (Trifolium repens) in Oklahoma. Plant Dis 96:1705–1706

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Phytopathology 86:1186–1192 Tosic M, Sutic D, Pesic Z (1980) Transmission of Tobacco mosaic virus through pepper (Capsicum annuum L. seed). Phytopathol Z 97:10–13 Truta AAC, Souza ARR, Nascimento AVS, Pereira RC, Pinto CMF, Brommonschenkel SH, Carvalho MG, Zerbini FM (2004) Identidade e propriedades de isolados de Potyvirus provenientes de Capsicum spp. Fitopatol Bras 29:160–168 Tsai WS, Shih SL, Green SK, Rauf A, Hidayat SH, Jan F-J (2006) Molecular characterization of Pepper yellow leaf curl Indonesia virus in leaf curl and yellowing diseased tomato and pepper in Indonesia. Plant Dis 90:247 Tsai WS, Huang YC, Zhang DY, Reddy K, Hidayat SH, Srithongchai W, Green SK, Jan F-J (2008) Molecular characterization of the CP gene and 30 UTR of Chilli veinal mottle virus from South and Southeast Asia. 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Phytopathology 105(3):388–398 Wetter C, Conti M, Altschuh D, Tabillion R, Van Regenmortel MHV (1984) Pepper mild mottle virus, a tobamovirus infecting pepper cultivars in Sicily. Phytopathology 74(4):405–410 Wetter C, Dore I, Bernard M (1987) Bell pepper mottle virus, a distinct tobamovirus infecting pepper. J Phytopathol 119:333–344 White KA (2011) Tombusvirus. Tombusviridae. The Springer index of viruses. Springer, New York, pp 1917–1926. https://doi.org/10.1007/978-0-387-95919-1_314 Wintermantel WM, Hladky LL (2013) Complete genome sequence and biological characterization of Moroccan pepper virus (MPV) and reclassification of Lettuce necrotic stunt virus as MPV. Phytopathology 103(5):501–508 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. 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Petria 9:342–343 Zaitlin M (2011) Tobamovirus. In: Tidona C, Darai G (eds) The Springer index of Viruses. Springer, New York. https:// doi.org/10.1007/978-0-387-95919-1 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zhang SB, Zhao ZB, Zhang DY, Liu Y, Zhang SB, Zhang DY, Liu Y, Luo XW, Liu J, Wu LF, Peng J (2015a) First teport of Pepper vein yellows virus infecting red pepper in mainland China. Plant Dis 99:1190 Zhang S, Zhang D, Liu Y, Liu J, Zhao Z, Luo X, Peng J (2015b) First report of chili pepper (Capsicum annuum) as a natural host plant for Dahlia mosaic virus. Plant Dis 99:898 Zhang SB, Liu J, Zhao ZB, Zheng LM, Zhang DY, Liu Y, Du J, Peng J, Yan F, Li F, Xie Y, Cheng ZB (2016a) First report of pepper (Capsicum annuum L.) as a natural host plant for Youcai mosaic virus. 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Capsicum annuum var. aviculare (Chiltepin)

431

Zhou X (2013) Advances in understanding Begomovirus satellites. Annu Rev Phytopathol 51:357–381 Zindovic J, Bulajic A, Krstic B, Ciuffo M, Margaria P, Turina M (2011) First report of Tomato spotted wilt virus on pepper in Montenegro. Plant Dis 95:882 Zitter TA (1972) Naturally occurring pepper virus strains in South Florida. Plant Dis Reptr 56:586–590

Capsicum annuum var. aviculare (Chiltepin) Family: Solanaceae

Chiltepin yellow mosaic virus Taxonomic position Genus: Tymovirus

(ChiYMV)

Family: Tymoviridae

Geographical distribution ChiYMV infection was isolated from plants of Capsicum annuum var. aviculare (Chiltepin) in Tula, Mexico (Pagan et al. 2010). Symptoms and host(s) The virus-infected chiltepin plants exhibit yellow mosaic symptoms in leaves. Host range is almost entirely limited to the Solanaceae, with systemic infections (mostly mosaic or yellow mosaic symptoms) produced in Capsicum annuum, Capsicum chinense, Capsicum frutescens, Datura stramonium, Nicotiana benthamiana, N. clevelandii, N. glutinosa, N. tabacum, N. rustica, Physalis floridana, and Solanum lycopersicum. Asymptomatic systemic infections were observed in Gomphrena globosa (Amaranthaceae) (Guy et al. 1984). Transmission The virus is transmitted by beetle vectors in a semi-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, isometric capsid with T = 3 icosahedral symmetry, about 30 nm in diameter. The genome consists of a single molecule of positive-sense ssRNA of 6517 nt (FN563123 = NC_014127). The 30 terminus has a tRNA-like structure (Martelli et al. 2002; Pagan et al. 2010).

References Guy PL, Dale JL, Adena MA, Gibbs AJ (1984) A taxonomic study of the host range of tymoviruses. Plant Pathol 33:337–346 Martelli GP, Sabanadzovic S, Abou-Ghanem Sabanadzovic N, Edwards MC, Dreher T (2002) The family Tymoviridae. Arch Virol 147(9):1837–1846 Pagan I, Betancourt M, de Miguel J, Pinero D, Fraile A, Garcia-Arenal F (2010) Genomic and biological characterization of Chiltepin yellow mosaic virus, a new tymovirus infecting Capsicum annuum var. aviculare in Mexico. Arch Virol 155:675–684

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432

Capsicum baccatum cv. Pendulum (Peruvian yellow chile pepper)

Capsicum baccatum cv. Pendulum (Peruvian yellow chile pepper) Family: Solanaceae

Spice crop

Pepper leafroll virus Taxonomic position Genus: Begomovirus

(PepLRV)

Family: Geminiviridae

Geographical distribution PepLRV infection in plants of Capsicum baccatum var. pendulum was reported from Peru (MartinezAyala et al. 2014). Symptoms and host(s) The virus-infected Peruvian yellow chile pepper plants show leaf roll symptoms. Natural occurrence of this virus is also reported on tomato, Nicandra physalodes, and common bean from Peru. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2568 nt (KC769819) (Martinez-Ayala et al. 2014; Brown et al. 2015; Zerbini et al. 2017).

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Capsicum baccatum was reported from Israel (Polston et al. 2006). The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The virus is transmissible by grafting but not transmissible by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

References Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Martinez-Ayala A, Sanchez-Campos S, Caceres F, Aragon-Caballero L, Navas-Castillo J, Moriones E (2014) Characterisation and genetic diversity of Pepper leaf roll virus, a new bipartite begomovirus infecting pepper, bean and tomato in Peru. Ann Appl Biol 164:62–72 Polston JE, Cohen L, Sherwood TA, Ben-Joseph R, Lapidot M (2006) Capsicum species: symptomless hosts and reservoirs of Tomato yellow leaf curl virus. Phytopathology 96:447–452 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Capsicum chinense (Yellow lantern chili)

433

Capsicum chinense (Yellow lantern chili) Family: Solanaceae

Vegetable

Capsicum chlorosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(CaCV)

Family: Tospoviridae

CaCV infection in plants of Capsicum chinense was reported from Australia (McMichael et al. 2002). The virus-infected plants exhibit stunting, leaf chlorosis, leaf distortion, and ringspot symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sapinoculation. For more details of CaCV, refer to Capsicum annuum.

Chilli ringspot virus Taxonomic position Genus: Potyvirus

(ChiRSV)

Family: Potyviridae

Geographical distribution ChiRSV infection in plants of Capsicum chinense was originally reported from Vietnam (Ha et al. 2008). The virus spreads in Hainan Province, China (Wang et al. 2012). Symptoms and host(s) The Hainan isolate of ChiRSV induces vein-banding and crinkle symptoms in yellow lantern chilli but not ringspot as described from the isolate reported from Vietnam. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically saptransmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome is a single molecule of positive-sense single-stranded RNA comprised of 9595 nt (JN008909 = NC_016044) excluding the 30 -terminal poly (A) tail and containing a large open reading frame of 9240 nt encoding a large polyprotein of 3079 amino acids with predicted Mr. of 349.1 kDa (DQ925438, DQ925439) (Gong et al. 2011; Wylie et al. 2017).

Chilli veinal mottle virus Synonyms Pepper vein banding virus

(ChiVMV)

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434

Taxonomic position Genus: Potyvirus

Capsicum chinense (Yellow lantern chili)

Family: Potyviridae

ChiVMV infection in plants of Capsicum chinense was reported from South and Southeast Asia and China (Wang et al. 2006; Banerjee et al. 2014). The virus-infected yellow lantern chilli plants exhibit dark green vein-banding mosaic and distorted leaf symptoms. The virus is transmitted by aphid vectors Aphis craccivora, A. gossypii, and Myzus persicae in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ChiVMV, refer to Capsicum annuum.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Capsicum chinense var. scotch bonnet was reported from Florida (USA) (Babu et al. 2014). The virus-infected yellow lantern chilli plants exhibit mosaic and yellow mottling symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Pepper vein yellows virus 1

(PeVYV-1)

Synonyms Pepper vein yellows virus (PeVYV); Pepper yellows virus

Taxonomic position Genus: Polerovirus

Family: Luteoviridae

PeVYV-1 infection in plants of Capsicum chinense was reported from Ivory Coast (Bolou Bi et al. 2015). The virus-infected plants exhibit symptoms of leaf yellowing and rolling as well as stunting. The virus is transmitted by aphid vectors in a circulative and non-propagative manner. For more details of PeVYV-1, refer to Capsicum annuum.

Tobacco mild green mosaic virus Taxonomic position Genus: Tobamovirus

(TMGMV)

Family: Virgaviridae

TMGMV was reported to infect Capsicum chinense in Venezuela (Cordoba et al. 2006). The virusinfected yellow lantern chilli plants exhibit symptoms of curling and bubbling of leaves. No insect vector is reported for this virus. The virus is transmissible by mechanical sap-inoculation, and has a wide host range. The virus is transmissible by contact between plants. The virus is transmissible by grafting and also through seed (Cordoba et al. 2006). For more details of TMGMV, refer to Nicotiana tabacum.

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Tomato chlorotic leaf distortion virus Taxonomic position Genus: Begomovirus

(ToClLDV)

Family: Geminiviridae

ToClLDV infection in plants of Capsicum chinense was reported from Venezuela (Zambrano et al. 2011). The virus-infected yellow lantern chilli plants exhibit leaf distortion and chlorotic mottle symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative nonpropagative manner. For more details of ToClLDV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Capsicum chinense was reported from Brazil and Florida (Boiteux and Nagata 1993; Momol et al. 2000). The virus-infected yellow lantern chilli plants exhibit symptoms of leaf necrosis, fruit drop, necrotic stem lesions, and stunting. Fruit symptoms included chlorotic and necrotic spotting and distinct ring pattern and distortion. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Capsicum chinense was reported from Israel (Polston et al. 2006). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

References Babu B, Dankers H, Paret ML (2014) First report of Cucumber mosaic virus associated with Capsicum chinense var. scotch bonnet in Florida. Plant Dis 98:1016 Banerjee A, Dutta R, Roy S, Ngachan SV (2014) First report of Chilli veinal mottle virus in Naga chilli (Capsicum chinense) in Meghalaya, India. Virus Dis 25:142–143 Boiteux LS, Nagata T (1993) Susceptibility of Capsicum chinense PI159236 to Tomato spotted wilt virus isolates in Brazil. Plant Dis 77:210 Bolou Bi BA, Moury B, Abo K Jr, Kakou D, Girardot G, Kouassi NP, Kouadio EJN, Kouakou BSM, Kone D (2015) First report of Pepper vein yellows virus in field-grown pepper in Ivory Coast. J Plant Pathol 97:S75 Cordoba MC, Garcia-Randez A, Montano N, Jorda C (2006) First report of Tobacco mild green mosaic virus in Capsicum chinense in Venezuela. Plant Dis 90:1108 Gong D, Wang JH, Lin ZS, Zhang SY, Zhang YL, Yu NT, Xiong Z, Liu ZX (2011) Genomic sequencing and analysis of Chilli ringspot virus, a novel Potyvirus. Virus Genes 43:439–444 Ha C, Revill P, Harding RM, Vu M, Dale JL (2008) Identification and sequence analysis of Potyviruses infecting crops in Vietnam. Arch Virol 153(1):45–60 McMichael LA, Persley DM, Thomas JE (2002) A new tospovirus serogroup IV species infecting capsicum and tomato in Queensland, Australia. Australas Plant Pathol 31:231–239

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Capsicum pubescens (Rukutu, Ruqutu, Rocoto)

Momol MT, Pappu HR, Dankers W, Rich JR, Olson SM (2000) First report of Tomato spotted wilt virus in habanero and tabasco peppers in Florida. Plant Dis 84:1154 Polston JE, Cohen L, Sherwood TA, Ben-Joseph R, Lapidot M (2006) Capsicum species: Symptomless hosts and reservoirs of Tomato yellow leaf curl virus. Phytopathology 96:447–452 Wang J, Liu Z, Niu S, Peng M, Wang D, Weng Z, Xiong Z (2006) Natural occurrence of Chilli veinal mottle virus on Capsicum chinense in China. Plant Dis 90:377 Wang J-H, Zhang S-Y, Gong D, Wu Y-P, Zhang Y-L, Yu N-T, Liu Z-X, Xiong Z (2012) First report of Chilli ringspot virus on Chili Pepper in China. Plant Dis 96:462 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Zambrano K, Geraud-Pouey F, Chirinos D, Romay G, Marys E (2011) Tomato chlorotic leaf distortion virus, a new bipartite begomovirus infecting Solanum lycopersicum and Capsicum chinense in Venezuela. Arch Virol 156: 2263–2266.

Capsicum pubescens (Rukutu, Ruqutu, Rocoto) Family: Solanaceae

Vegetable

Ecuadorian rocoto virus

(EcRV)

Taxonomic position EcRV is a tentative member of the genus Potyvirus and family Potyviridae. Geographical distribution EcRV infection in plants of Capsicum pubescens was reported from Riobamba (Ecuador) (Janzac et al. 2008). Symptoms and host(s) The virus-infected rocoto plants exhibit mild or severe mosaic symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. Virion properties and genome The virions are flexuous and 700–900 nm long. The genome is a positive-sense single-stranded RNA. A partial genome sequence of 2951 nt is available (EU495234).

References Janzac B, Fabre MF, Palloix A, Moury B (2008) Characterization of a new Potyvirus infecting pepper crops in Ecuador. Arch Virol 153(8):1543–1548

Cardamine spp. Family: Brassicaceae

Weed host

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

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BWYV infection in plants of Cardamine spp. was reported from Australia (Guy 1987; Ellis 1992). The virus-infected Cardamine plants exhibit reddening of the lower leaves, interveinal chlorosis, and stunting. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of BWYV, refer to Beta vulgaris.

Cardamine chlorotic fleck virus Taxonomic position Genus: Betacarmovirus

C

(CCFV)

Family: Tombusviridae

Geographical distribution CCFV infection was first reported in Cardamine lilacina plants from Australia by Skotnicki et al. (1992, 1993). The virus spreads in Australia (Skotnicki and Mo 1996). Symptoms and host(s) The virus-infected Cardamine plants exhibit chlorotic leaf flecking symptoms. Transmission The virus is transmitted by mechanical sap-inoculation. Virion properties and genome The virions are isometric 30 nm in diameter and have T = 3 icosahedral symmetry. The genome consists of a single molecule of linear positive-sense ssRNA of 4041 nt (L16015 = NC_001600) (Skotnicki et al. 1993).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Cardamine scutata was reported from Japan (Okuda et al. 2010). The virusinfected Cardamine plants exhibit necrotic spot symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Cardamine oligosperma was reported from British Columbia and the USA (Bitterlich and MacDonald 1993; Groves et al. 2002). The virus-infected Cardamine plants exhibit yellow spots and ringspot symptoms. The virus is transmitted by a thrips vector, Frankliniella fusca, in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of host plants. For more details of TSWV, refer to Solanum lycopersicum.

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Turnip yellow mosaic virus

(TYMV)

Synonyms Cardamine yellow mosaic virus Taxonomic position Genus: Tymovirus

Family: Tymoviridae

TYMV infection in plants of Cardamine lilacina was reported from Australia (Guy and Gibbs 1981, 1985; Gibbs et al. 1986; Skotnicki et al. 1996). The virus-infected Cardamine plants exhibit bright veinclearing and yellow mosaic symptoms. The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sap-inoculation, and by grafting. For more details of TYMV, refer to Brassica rapa.

References Bitterlich I, MacDonald LS (1993) The prevalence of Tomato spotted wilt virus in weeds and crops in southwestern British Columbia. Can Plant Dis Surv 73(2):137–142 Ellis PJ (1992) Weed hosts of Beet western yellows virus and Potato leafroll virus in British Columbia. Plant Dis 76:1137–1139 Gibbs AJ, Blok J, Coates DJ, Guy PL, Mackenzie A, Pigram N (1986) Turnip yellow mosaic virus in an endemic Australian alpine Cardamine. In: Barlow BA (ed) Flora and fauna of alpine Australasia: ages and origins. CSIRO, Australia, pp 289–299 Groves RL, Walgenbach JF, Mayor JW, Kennedy GG (2002) The role of weed hosts and tobacco thrips, Frankliniella fusca, in the epidemiology of Tomato spotted wilt virus. Plant Dis 86:573–582 Guy PL (1987) Beet western yellows virus infection of Cardamine spp. and Cheesemania radicata. Australas Plant Pathol 16:43–44 Guy PL, Gibbs AJ (1981) A Tymovirus of Cardamine sp. from alpine Australia. Australas Plant Pathol 10:12–13 Guy PL, Gibbs AJ (1985) Further studies on Turnip yellow mosaic tymovirus isolates from an endemic Australian Cardamine. Plant Pathol 34:532–544 Okuda M, Fuji S, Okuda S, Sako K, Iwanami T (2010) Evaluation of the potential of thirty two weed species as infection sources of Impatiens necrotic spot virus. J Plant Pathol 92:357–361 Skotnicki ML, Mo J-Q (1996) Detection of Cardamine chlorotic fleck carmovirus in soil by the polymerase chain reaction. Australas Plant Pathol 25:18–23 Skotnicki ML, Mackenzie AM, Torronen M, Brunt AA, Gibbs AJ (1992) Cardamine chlorotic fleck virus, a new Carmovirus from the Australian Alps. Australas Plant Pathol 21:120–122 Skotnicki ML, Mackenzie AM, Torronen M, Gibbs AJ (1993) The genomic sequence of Cardamine chlorotic fleck carmovirus. J Gen Virol 74:1933–1937 Skotnicki ML, Mackenzie AM, Gibbs AJ (1996) Genetic variation of Turnip yellow mosaic tymovirus within single plants of Cardamine lilacina. Australas Plant Pathol 25:249–254

Carica papaya (Papaya) Family: Caricaceae

Edible fruit crop

Ageratum yellow vein virus Taxonomic position Genus: Begomovirus

(AYVV)

Family: Geminiviridae

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AYVV infection in plants of Carica papaya was reported from China and India (Shahid et al. 2013; Shen et al. 2014a). The virus-infected papaya plants exhibit downward curling of the leaves and yellow vein symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative nonpropagative manner. The virus is not transmissible by mechanical sap-inoculation, and is also not by contact between plants. For more details of AYVV, refer to Ageratum spp.

C Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

(AMV)

Family: Bromoviridae

AMV infection in plants of Carica papaya was reported from Brazil (Moreira et al. 2010). The virusinfected papaya plants exhibited severe yellow leaf mosaic, leaf distortion, and systemic necrosis symptoms. Even though AMV is not currently considered to be a major threat to the papaya crop, papaya plants in Brazil and elsewhere may be endangered if favorable conditions for virus dissemination and infection are established. The virus is aphid transmitted in a non-persistent manner, and also through mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Chickpea chlorotic dwarf virus Taxonomic position Genus: Mastrevirus

(CpCDV)

Family: Geminiviridae

CpCDV infection in plants of Carica papaya was reported from Nigeria (Fiallo-Olive et al. 2017; Ouattara et al. 2017). The virus-infected papaya plants exhibit stunting, leaf deformation, curling, and yellowing symptoms. The virus is transmitted by leafhopper vectors in a persistent, circulative, and non-propagative manner. The virus is not transmissible by mechanical inoculation. For more details of CpCDV, refer to Cicer arietinum.

Cotton leaf curl Gezira virus Taxonomic position Genus: Begomovirus

(CLCuGeV)

Family: Geminiviridae

CLCuGev infection in plants of Carica papaya was reported from Oman (Khan et al. 2012). The virusinfected papaya plants exhibit symptoms of leaves with severe curling, vein darkening, and vein thickening. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative nonpropagative manner. The virus is not transmissible by mechanical sap-inoculation or by contact between plants. For more details of CLCuGeV, refer to Gossypium spp.

Cotton leaf curl Multan virus Taxonomic position Genus: Begomovirus

(CLCuMuV)

Family: Geminiviridae

440

Carica papaya (Papaya)

CLCuMuV infection in plants of Carica papaya was reported from India (Sinha et al. 2013). The virusinfected papaya plants exhibit leaf curling symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of CLCuMuV, refer to Gossypium spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Carica papaya was first reported from Puerto Rico by Adsuar (1956). This virus was also reported from India (Kiranmai et al. 1998; Tennant et al. 2007). This virus-infected papaya plants exhibit mosaic, leaf distortion, and philiform symptoms. The virus-infected papaya plants are stunted and produce very few small and deformed fruits. The virus is transmitted by a number of aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Euphorbia leaf curl virus Taxonomic position Genus: Begomovirus

(EuLCuV)

Family: Geminiviridae

EuLCuV infection in plants of Carica papaya was reported from Korea (Kil et al. 2016). The virusinfected papaya plants exhibit symptoms of mild yellow spotting and curling of leaves. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of EuLCuV, refer to Euphorbia pulcherrima.

Malvastrum leaf curl virus Taxonomic position Genus: Begomovirus

(MaLCuV)

Family: Geminiviridae

MaLCuV infection in plants of Carica papaya was reported from China (Wu and Zhou 2006). The virus-infected papaya plants exhibit leaf curl symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of MaLCuV, refer to Malvastrum spp.

Moroccan watermelon mosaic virus Taxonomic position Genus: Potyvirus

(MWMV)

Family: Potyviridae

MWMV infection in plants of Carica papaya was reported from the Democratic Republic of Congo (Arocha et al. 2008). The virus-infected papaya plants exhibit yellow mosaic, deformation, and

Carica papaya (Papaya)

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shoestring symptoms on leaves. Oily streaks develop on the petiole and ringspot symptoms on fruits. The virus is transmitted by aphid vectors Myzus persicae and Aphis gossypii in a non-persistent manner, and also by mechanical sap-inoculation only to species belonging to the family Cucurbitaceae. For more details of MWMV, refer to Citrullus lanatus.

Okra enation leaf curl virus Taxonomic position Genus: Begomovirus

(OELCuV)

Family: Geminiviridae

OELCuV infection in plants of Carica papaya was reported from Iran (Bananej et al. 2016). The virusinfected papaya plants exhibit severe leaf curling and vein swelling symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of OELCuV, refer to Abelmoschus esculentus.

Papaya leaf crumple virus Taxonomic position Genus: Begomovirus

(PaLCrV)

Family: Geminiviridae

Geographical distribution PaLCrV infection in plants of Carica papaya plants was reported from India (Singh-Pant et al. 2012). Symptoms and host(s) The virus-infected papaya plants exhibit leaf crumple symptoms. The virus has been isolated from papaya (Carica papaya), soybean (Glycine max), Catharanthus roseus, Andrographis paniculata, and Solanum nigrum. Transmission The transmission of PaLCrV has not been investigated. It is likely that, in common with other begomoviruses, PaLCrV is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of PaLCrV has not been investigated. In common with all geminiviruses, the virions of PaLCrV are likely geminate (twinned quasi-icosahedra). PaLCrV is a typical old-world monopartite begomovirus with a genome that consists of a single circular molecule of single-stranded DNA of ~2736 nt (HM140367 = NC_014707; HE580236, HM140368, HM140369) (Briddon 2001; Singh-Pant et al. 2012; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes of PaLCrV encode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated for PaLCrV. PaLCrV associates with betasatellites (Singh-Pant et al. 2012).

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442

Carica papaya (Papaya)

Papaya leaf curl China virus Taxonomic position Genus: Begomovirus

(PaLCuCNV)

Family: Geminiviridae

Geographical distribution PaLCuCNV infection in plants of Carica papaya was reported from China and Vietnam (Wang et al. 2004; Cai et al. 2005; Tennant et al. 2007; Ha et al. 2008). Symptoms and host(s) The virus-infected papaya plants exhibit leaf curl symptoms. The virus has been identified infecting papaya (Carica papaya), Ageratum conyzoides, tomato (Solanum lycopersicum), Corchoropsis timentosa, Eclipta prostrata, tobacco (Nicotiana tabacum), and Siegesbeckia orientalis. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Transmission of PaLCuCNV was studied using whitefly genotypes, Middle East-Asia Minor 1 (MEAM1), Mediterranean (MED), Asia 1, and Asia II 7 that showed difference in levels of transmission efficiency. In tomato, MEAM1 showed a higher efficiency of transmission of PaLCuCNV than MED, Asia 1, and Asia II (Guo et al. 2015). Virion properties and genome The structure of the virions of PaLCuCNV has not been investigated. In common with all geminiviruses, the virions of PaLCuCNV are likely geminate (twinned quasi-icosahedra). PaLCuCNV is a typical old-world monopartite begomovirus with a genome that consists of a single circular molecule of single-stranded DNA of ~2751 nt (AJ558124 = NC_005321; AJ558116, AJ558117, AJ558123, AJ558125, AJ704604, AJ811439, AJ811914, AJ876548, AM691552, AM691553, AM691554, DQ641700, EU874386, FN256260, FN256261, FN297834, FN434083, GQ338766, GQ373254, GQ870288, HG003651, HQ162268, HQ162269, JF682837, JX128101, JX128102, JX294075, JX555979, KC857511, KC878473, KC878474). Papaya leaf curl China betasatellite DNA molecule is associated with PaLCuCNV, and consists of 1350 nt (KJ642219) (Briddon 2001; Wang et al. 2004; Cai et al. 2005; Ha et al. 2008; Zhou 2013; Brown et al. 2015; Guo et al. 2015; Zerbini et al. 2017). The characterized genomes of PaLCuCNV encode the six genes typically encoded by monopartite begomoviruses originating from the Old World: two in the virion-sense and four in the complementarysense. The expression and function of the genes have not been investigated for PaLCuCNV.

Papaya leaf curl Coimbatore virus

(PaLCCoV)

Taxonomic position PaLCCoV is a tentative member of the genus Begomovirus and family Geminiviridae. Geographical distribution PaLCCoV infection in plants of Carica papaya plants was reported from Andhra Pradesh, Karnataka, Tamil Nadu, Uttar Pradesh, and Delhi states of India (Krishna Reddy et al. 2010).

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Symptoms and host(s) The virus-infected papaya plants exhibit symptoms of leaf curl, vein-clearing, enations, and reduced leaf size. Transmission Vector transmission has not been investigated. The virus is expected to be transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The structure of PaLCCoV has not been investigated. In common with all geminiviruses, the virions of PaLCCoV are likely geminate (twinned icosahedra).

Papaya leaf curl Guandong virus Taxonomic position Genus: Begomovirus

(PaLCuGdV)

Family: Geminiviridae

Geographical distribution PaLCuGdV-isolate GT infecting plants of Carica papaya was detected and obtained from Guangzhou, Guangdong Province, China (Wang et al. 2004; Zhang et al. 2005; Tennant et al. 2007). Some isolates of the virus were previously called Malvastrum leaf curl Guangdong virus. Symptoms and host(s) The virus-infected papaya plants show typical leaf curl symptoms. The virus has been isolated from papaya (Carica papaya), Malvastrum coromandelianum, Euphorbia pulcherrima, tobacco (Nicotiana tabacum), Euphorbia pulcherrima, Blechum pyramidatum, and passionfruit hybrid (Passiflora edulis x Passiflora edulis). Transmission The transmission of PaLCuGdV has not been investigated. It is likely that, in common with other begomoviruses, PaLCuGdV is transmitted by the whitefly Bemisia tabaci in a circulative and nonpropagative manner. Virion properties and genome The structure of the virions of PaLCuGdV has not been investigated. In common with all geminiviruses, the virions of PaLCuGdV are likely geminate (twinned quasi-icosahedra). PaLCuGdV is a typical old-world monopartite begomovirus with a genome that consists of a single circular molecule of single-stranded DNA of ~2742 nt (AJ558122 = NC_005844; AM236779, AM236780, AM503104, AY650283, EF554783, FJ495184, FJ712189, FJ869907, JN703795, KC161184, KF446659, KF446660) (Briddon 2001; Zhang et al. 2005; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes of PaLCuGdV encode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated for PaLCuV.

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Papaya leaf curl virus Taxonomic position Genus: Begomovirus

Carica papaya (Papaya)

(PaLCuV)

Family: Geminiviridae

Geographical distribution PaLCuV was first reported in plants of Carica papaya from Puerto Rico as “curly leaf” (Cook 1931). The virus spreads in Puerto Rico, Guiana, the Philippines, Taiwan, Pakistan, Korea, and India (Nariani 1956; Verma and Prasad 1985/1986; Nadeem et al. 1997; Saxena et al. 1998; Chang et al. 2003; Zhang et al. 2005; Singh 2006; Usharani et al. 2013; Dubey et al. 2015; Byun et al. 2016). Symptoms and host(s) The virus-infected papaya plants exhibit symptoms of severe curling and crinkling, distortion of leaves, and reduction of petiole, internodes, and also the main shoot. The leaves are drastically reduced in size and show vein-clearing. Leaf margins are rolled downward and inward in the form of inverted cups. Leaves of infected plants turn dark green and become leathery and brittle. The petioles are twisted in a zigzag manner, and bend toward the main trunk; at times enations are produced on the lower surface of the leaves. The virus has been isolated from papaya (Carica papaya), tomato (Solanum lycopersicum), radish (Raphanus sativus), Cape gooseberry (Physalis peruviana), guar (Cyamopsis tetragonoloba), Croton bonplandianus, cotton (Gossypium hirsutum), Aster alpinus, Amaranthus cruentus, Nicotiana glutinosa, N. tabacum, Capsicum spp., Physalis peruviana, Rhynchosia capitata, Crotalaria juncea, and soybean (Glycine max). Transmission The transmission of PaLCuV has not been investigated. It is likely that, in common with other begomoviruses, PaLCuV is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible (Dubey et al. 2015). Virion properties and genome The structure of the virions of PaLCuV has not been investigated. In common with all geminiviruses, the virions of PaLCuV are likely geminate (twinned quasi-icosahedra). PaLCuV is a typical old-world monopartite begomovirus with a genome that consists of a single circular molecule of single-stranded DNA of ~2746 nt (Y15934 = NC_004147; FM955602, DQ629102) (Briddon 2001; Krishna Reddy et al. 2010; Brown et al. 2015; Zerbini et al. 2017). Papaya leaf curl betasatellite DNA molecule is associated with PaLCuV, and consists of 1372 nt (AY244706) is associated with PaLCuV (Zhou 2013). The characterized genomes of PaLCuV encode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated for PaLCuV.

Papaya leaf distortion mosaic virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

(PLDMV)

Carica papaya (Papaya)

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Geographical distribution PLDMV infection in plants of Carica papaya was first reported in Okinawa during 1954 (Kawano and Yanaha 1992). The virus spreads in China, Taiwan, and Japan (Kawano and Yanaha 1992; Maoka et al. 1996; HuoGen et al. 1997; Xiao et al. 1997; Bau et al. 2008; Shen et al. 2014b). Symptoms and host(s) The virus-infected papaya plants show vein-clearing and then mosaic and distortion of leaves, leaf narrowing and shoestringing symptoms, apex stunting in the infected plants, and water-soaked streaks may appear on the petioles and stems and dark green brown ringspot symptoms on fruits. Transmission The virus is transmitted by aphid vectors, viz., Aphis gossypii, A. citricola, A. nerii, A. clerodendri, A. crassivora, and Myzus persicae, in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 10,155 nt (BD171712 = NC_005028) (Maoka et al. 1996; Maoka and Hataya 2005; Revers and Garcia 2015; Wylie et al. 2017).

Papaya lethal yellowing virus Taxonomic position Genus: Sobemovirus

(PLYV)

Family: Solemoviridae

Geographical distribution PLYV infection in plants of Carica papaya was reported only from Brazil (Loreto et al. 1983; Silva et al. 1997; Teixeira et al. 1999; Tennant et al. 2007; Ramos et al. 2008; Lima et al. 2013). Symptoms and host(s) The virus-infected papaya plants exhibit symptoms beginning with a progressive yellowing of the upper leaves that eventually fall off; as the disease advances, the leaves curl and become chlorotic followed by wilting and senescence and subsequently die followed by the death of the entire plant. Greenish circular spots appear on the fruits that turn yellowish as the fruits ripen (Lima et al. 2001). Transmission The virus is readily sap-transmissible and found in the soil, but a vector has not been identified. The virus is transmissible by wounding and grafting. The virus is also transmissible through contaminated hands. No seed transmission was reported (Camarco et al. 1998; Saraiva et al. 2006). Virion properties and genome The virions are isometric, non-enveloped, and 30 nm in diameter. The genome is a polycistronic positive-sense single-stranded RNA (ssRNA) of 4145 nt (JX123318 = NC_018449). The genome contains two overlapping ORFs, ORF2a and ORF2b (Kitajima et al. 1992; Kitajima 1999; Nascimento et al. 2010; Pereira et al. 2012; Ling et al. 2013; Somera et al. 2015).

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Papaya meleira virus

Carica papaya (Papaya)

(PMeV)

Taxonomic position PMeV is a tentative member of the family Totiviridae. Geographical distribution PMeV infection in plants of Carica papaya was first observed in southern Bahia (Brazil), and local growers refer this disease as “meleira” (sticky disease) (Nakagawa et al. 1987). The virus spreads in Brazil, Indonesia, and Mexico (Rodrigues et al. 1989; Maciel-Zambolim et al. 2003; Araujo et al. 2007; Tennant et al. 2007; Perez-Brito et al. 2012; Daltro et al. 2014; Abreu et al. 2012, 2015; Sá Antunes et al. 2016). Symptoms and host(s) The virus-infected papaya plants exhibit symptoms of latex exudation in petioles, new leaves, and fruits; when the exudate oxidizes, it causes necrosis of the affected tissues. The latex of diseased plants loses viscosity and is easily dispersed. First symptoms appear on young leaves of 6-month-old plants; eventually they become distorted and develop a burnt appearance. Dark spots are present on the pulp of the fruit. The most distinct and typical symptom produced by Papaya meleira virus is the spontaneous exudation of translucent and watery latex of an infected fruit (Rodrigues et al. 1989, 2009a). It has been reported that the ‘meleira’ disease does not occur in plants singly infected with PMeV, but requires coinfection with a presumed umbravirus, Papaya meleira virus 2 (Sá Antunes et al. 2016). Transmission The virus is not mechanically transmitted, and the whitefly Trialeurodes variabilis failed to transmit this disease (Rodrigues et al. 2009b), although injection of partially purified extracts was infectious (Maciel-Zambolim et al. 2003). From Mexico, Tapia-Tussell et al. (2015) have reported that this disease is transmissible through papaya seed at the rate of c. 81%, suggesting PMeV-contaminated seed is the source of transmission. Virion properties and genome The virions are isometric particles, 50 nm in diameter. The genome is double-stranded RNA consists of 8768 nt (NC_028378) and contains two putative ORFs; the predicted ORF1 and ORF2 display similarity to capsid proteins and RdRp’s (Kitajima et al. 1993; Abreu et al. 2015; Sá Antunes et al. 2016).

Papaya meleira virus 2

(PMeV2)

Taxonomic position PMeV2 is currently unassigned, but is a presumed member of the genus Umbravirus, family Tombusviridae (Sá Antunes et al. 2016). Geographical distribution This virus was reported from Brazil affecting Carica papaya plants affected with “meleira” (sticky disease) (Sá Antunes et al. 2016). This disease occurs in Brazil, with similar symptoms observed in Mexico.

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Symptoms and host(s) Papaya plants infected with “meleira” (sticky disease), which probably requires both PMeV and PMeV2 (Sá Antunes et al. 2016), show exudation of liquid latex, leading to necrotic lesions at leaf margins and stickiness on the fruits. No other hosts are reported. Transmission The virus has no known particles of its own, and is probably transmitted by encapsidation within virions of PMeV, which is seed-transmitted in papaya (Tapia-Tussell et al. 2015). Transmission of the helper virus PMeV by injection has also been reported (Maciel-Zambolim et al. 2003). Virion properties and genome PMeV2 produces no coat protein or particles of its own, but in encapsidated by the associated PMeV; the genome is single-stranded RNA, and a partial sequence of 4515 nt is available (KT921785) (Sá Antunes et al. 2016).

Papaya mosaic virus Taxonomic position Genus: Potexvirus

(PapMV)

Family: Alphaflexiviridae

Geographical distribution PapMV infection in plants of Carica papaya was first reported by Conover (1962). The virus spreads in India, Pakistan, Srilanka, Bolivia, Peru, the USA, Jamaica, Mexico, Nigeria, and Venezuela (Lana 1980; Rajapakse and Herath 1981; Taya and Singh 1995; Noa-Carrazana and Silva-Rosales 2001; NoaCarrazana et al. 2006; Tennant et al. 2007). Symptoms and host(s) The virus-infected papaya plants show symptoms of vein-clearing or vein-banding and mosaic mottling of newly affected leaves. The older leaves show marginal curling and chlorotic spots. The lamina of the leaves is reduced and modified. The virus-infected plants are stunted. Symptoms are usually absent on petioles, stems, and fruits. Transmission The virus is transmitted by means not involving a vector (Cheema and Reddy 1985). The virus is mechanically sap-transmissible to plant species in 19 dicotyledon families. The virus is not seedtransmitted. Virion properties and genome The virions are flexuous filaments, non-enveloped with a clear modal length of 530 nm and 13 nm wide. The genome is single-stranded RNA of 6656 nt (D13957 = NC_001748) and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type, 18–27 kDa in size (Hiebert 1970; Bhaskar 1983; Lok and Abouhaidar 1986; Abouhaidar 1988; Sit et al. 1989; Adams et al. 2004).

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Papaya ringspot virus Taxonomic position Genus: Potyvirus

Carica papaya (Papaya)

(PRSV)

Family: Potyviridae

Geographical distribution PRSV infection in plants of Carica papaya was first reported from the Island of Oahu in 1938. The virus spreads worldwide including the Middle East and the South and Central American region, Europe, Australia, China, France, Germany, India, Iran, Italy, Mexico, Taiwan, the Philippines, Thailand, Japan, Africa, Okinawa, Sri Lanka, Bangladesh, Nigeria, Colombia, Brazil, Kenya, and the USA (Lana 1980; Wan and Conover 1983; Opina 1986; Prasad and Sarkar 1989; Teliz et al. 1991; Bateson et al. 1994; Gonsalves 1994; Dahal et al. 1997; HuoGen et al. 1997; Xiao et al. 1997; Perera et al. 1998; Kiritani and Su 1999; Marys et al. 2000; Natsuaki and Bajet 2001; Ndunguru and Rajababu 2002; CABI/EPPO 2003; Pourrahim et al. 2003; Jain et al. 2004; Noa-Carrazana et al. 2006; Diallo et al. 2007; Tennant et al. 2007; Chin et al. 2008; Diallo et al. 2008; Gonsalves et al. 2010; Dhanam et al. 2011; Usharani et al. 2013; Ombwara et al. 2014; Shen et al. 2014b; Martins et al. 2016; Cabrera Mederos et al. 2016; Zhao et al. 2016; Chittarath et al. 2017; Ortiz-Rojas and Chaves-Bedoya 2017). Symptoms and host(s) PRSV (P-strain) symptoms on papaya may vary in intensity according to the age at which the plant becomes infected and the strain of the virus. Leaf symptoms are characterized by intense yellow mosaic and leaf distortion. The leaf laminae are markedly reduced in size and may develop a shoestring appearance. Oily streaks on the stem and petioles of the leaves are frequently observed on diseased plants. Several circular or concentric water-soaked lesions and necrotic rings with a solid central spot are almost always present on fruits. Symptoms are severe in cool weather (Thomas and Dodman 1993; Husain 1997; Jain et al. 1998; Davis et al. 2005). PRSV-W strain (Watermelon mosaic virus-1) which infects cucurbits is also widespread wherever papaya is grown. Inoue-Nagata et al. (2007), Gude et al. (2008), and Yeh et al. (1984) have reported the comparative studies between PRSV-P and PRSV-W strains. The host range of PRSV (P-strain) is limited to plants in the families Caricaceae, Cucurbitaceae, and Chenopodiaceae. Chenopodium amaranticolor and C. quinoa are reported as local lesion hosts of PRSV-P. In addition to the cultivated Carica papaya additional species including C. goudotiana, C. horovitziana, C. microcarpa, C. monoica, C. parviflora, and C. quercifolia are also susceptible to PRSV-P (Thomas and Dodman 1993; Maoka et al. 1995; Tripathi et al. 2008). Transmission The virus is transmitted by more than 20 aphid vector species including Myzus persicae, Aphis coreopsidis, A. craccivora, A. fabae, A. gossypii, Toxoptera citricida, etc., in a non-persistent manner (Lana 1980; Wang 1981; Hwang and Hsieh 1984; Taya and Singh 1997; Sharma et al. 2005; CortezMadrigal and Mora-Aguilera 2007; Kalleshwaraswamy and Kumar 2008; Martins et al. 2016). The virus is transmissible by mechanical sap-inoculation to several herbaceous host species. Bayot et al. (Bayot et al. 1990) have reported very low (0.15%) seed transmission of PRSV in papaya from the Philippines. Virion properties and genome The virions are flexuous, filaments, and non-enveloped, with a clear modal length of 760–800 nm and 12 nm width (De La Rosa and Lastra 1983). The genome consists of a single molecule of linear, positive-sense ssRNA of 10,326 nt (X67673 = NC_001785). The genome is monocistronic expressing

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a large polyprotein, which is subsequently processed into individual functional proteins. The virus is encapsidated by coat protein of molecular weight of 32–36 kD. It has a single large open reading frame that starts at nucleotide position 86–88 and ends at the positions 10118–10120 encoding a polyprotein of 3344 amino acids (Yeh and Gonsalves 1985; Quemada et al. 1990; Bateson and Dale 1992; Yeh et al. 1992; Jain et al. 1998; Kunkalikar and Byadgi 2004; Parameswari et al. 2007; Tripathi et al. 2008; Sharma et al. 2014; Revers and Garcia 2015; Wylie et al. 2017).

Papaya virus Q

(PpVQ)

Taxonomic position PpVQ is currently unassigned, but has an Umbravirus-like genome, and thus probably belongs in the family Tombusviridae (Quito-Avila et al. 2015). Geographical distribution PpVQ has been reported only from Ecuador, most commonly in plants coinfected with papaya ringspot virus (PRSV) (Quito-Avila et al. 2015; Cornejo-Franco et al. 2018). Symptoms and host(s) Papaya plants singly infected with PpVQ showed no obvious symptoms, and plants dually infected with PRSV showed no symptoms different from those caused by PRSV alone (Cornejo-Franco et al. 2018). Symptomatic plants of babaco (Vasconcellea x heilbornii, syn. Carica pentagona) were also found to be infected with a virus with 77% nt identity to PpVQ, but the symptoms were not described (Cornejo-Franco et al. 2018). Transmission PpVQ was not found to be either transmitted through seed or by mechanical inoculation, but three of ten plants exposed to whiteflies (Bemisia tabaci) collected from PpVQ-infected plants tested positive for PpVQ at 90 days post exposure (Cornejo-Franco et al. 2018); in the field, PpVQ incidence increased during the growing season, suggesting transmission by an aerial vector (Quito-Avila et al. 2015). PpVQ was not transmitted by aphids, nor by mechanical inoculation from plants coinfected with PRSV (Quito-Avila et al. 2015). Vrion properties and genome No virions have been identified. The genome is single-stranded RNA of c.4.5 kb, but a partial sequence of PpVQ (as Papaya umbra virus Ec_12; KP165407) lacks the 30 ; region, possibly due to RNA secondary structures interfering with priming of cDNA (Quito-Avila et al. 2015).

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Carica papaya was reported from Nigeria and the Rio Grande Valley (Texas) (Mclean and Olson 1962; Lambe 1963; Lana 1980; Tennant et al. 2007). The virus-infected papaya plants show wilting symptoms and develop terminal necrosis before they are killed. The virus is transmitted by a nematode vector Xiphinema americanum in a non-persistent manner, and also by

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mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Carica papaya was reported from Lucknow (India) (Raj et al. 2008). The virus-infected papaya plants showed severe downward leaf curling, swelling of veins, twisting and reduction of petioles, and stunted growth of the whole plant which bore a few small, distorted fruit. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The disease was successfully transmitted by B. tabaci from naturally infected papaya to healthy seedlings of papaya, tobacco, tomato, and chilli. All the inoculated test species displayed typical leaf curl symptoms. The virus is graft-transmissible. For more details of ToLCNDV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV was first observed in plants of Carica papaya on the island of “Kauai” (Hawaii) in 1962 (Trujillo and Gonsalves 1967). The virus spreads in the Hawaiian Islands, the USA, and Brazil (Gonsalves and Trujillo 1986; Rezende and Costa 1987; Tennant et al. 2007). The virus-infected papaya plants exhibit severe chlorosis and necrosis of apical leaves. Plants defoliate prematurely, and young infected plants show pronounced bending of the stem apex. Dark green water-soaked areas are found in petioles and stems with green or gray circular spots on young fruits, and mature green fruits exhibit dark green ringspots. The virus is transmitted by thrips vectors, Thrips tabaci, Frankliniella schulzei, F. fusca, F. occidentalis, and Scirtothrips dorsalis, in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

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FFTC Book Series No. 33, 1986. pp 157–168 Ortiz-Rojas LY, Chaves-Bedoya G (2017) Molecular characterization of two Papaya ringspot virus isolates that cause devastating symptoms in Norte de Santander, Colombia. Eur J Plant Pathol 148:883–894 Ouattara A, Tiendrebeogo F, Lefeuvre P, Hoareau M, Claverie S, Traore EV, Barro N, Traore O, Varsani A, Lett J-M (2017) New strains of Chickpea chlorotic dwarf virus discovered on diseased papaya and tomato plants in Burkina Faso. Arch Virol 162:1791–1794 Parameswari B, Mangrauthia SK, Praveen S, Jain RK (2007) Complete genome sequence of an isolate of Papaya ringspot virus from India. Arch Virol 152:843–845 Pereira AJ, Alfenas-Zerbini P, Cascardo RS, Andrade EC, Murilo Zerbini F (2012) Analysis of the full-length genome sequence of Papaya lethal yellowing virus (PLYV), determined by deep sequencing, confirms its classification in the genus Sobemovirus. Arch Virol 157(10):2009–2011 Perera WGS, Suetsugu T, Saito N (1998) Occurrence of Papaya ring spot potyvirus strain P in Sri Lanka. JICA/NPQS Occas Pap 1:14 Perez-Brito D, Tapia-Tussell R, Cortes-Velazquez A, Quijano-Ramayo A, Nexticapan-Garcez A, Martin-Mex R (2012) First report of Papaya meleira virus (PMeV) in Mexico. Afr J Biotechnol 11:13564–13570 Pourrahim R, Farzadfar S, Golnaraghi AR, Shahraeen N (2003) First report of Papaya ringspot virus on papaya in Iran. Plant Dis 87(9):1148 Prasad SM, Sarkar DP (1989) Some ecological studies on Papaya ringspot virus in Ranchi. Indian J Virol 5:118–122

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Carica papaya (Papaya)

Quemada H, L'Hostis B, Gonsalves D, Reardon IM, Heinrikson R, Hiebert EL, Sieu LC, Slightom JL (1990) The nucleotide sequences of the 3-terminal regions of papaya ringspot virus strains W and P. J Gen Virol 71(1):203–210 Quito-Avila DF, Alvarez RA, Ibarra MA, Martin RR (2015) Detection and partial genome sequence of a new umbra-like virus of papaya discovered in Ecuador. Eur J Plant Pathol 143:199–204 Raj SK, Snehi SK, Khan MS, Singh R, Khan AA (2008) Molecular evidence for association of Tomato leaf curl New Delhi virus with leaf curl disease of papaya (Carica papaya L.) in India. Aust Plant Dis Notes 3(1):152–155 Rajapakse RH, Herath HM (1981) Host susceptibility of the Papaya mosaic-virus in Sri-Lanka. Beitr Trop Landwirt 19:429–432 Ramos NF, Nascimento AKQ, Goncalves MFB, Lima JAA (2008) Presenca dos virus da mancha anelar e do amarelo letal em frutos de mamoeiro comercializados. Trop Plant Pathol 33:449–452 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Rezende J, Costa A (1987) Four viruses infecting papaya plants experimentally. Fitopatol Bras 12:63–65 Rodrigues CH, Alves FL, Marin SLD (1989) Occurrence and symptoms of “Meleira” in papaya (Carica papaya L.) in the Espirito Santo State, Brazil. Fitopatol Bras 14:118 Rodrigues SP, da Cunha M, Ventura JA, Fernandes PMB (2009a) Effects of the Papaya meleira virus on papaya latex structure and composition. Plant Cell Rep 28:861–871 Rodrigues SP, Andrade JS, Ventura JA, Lindsey GG, Fernandes PMB (2009b) Papaya meleira virus is neither transmitted by infection at wound sites nor by the whitefly Trialeurodes variabilis. J Plant Pathol 91:87–91 Sá Antunes TF, Amaral RJ, Ventura JA, Godinho MT, Amaral JG, Souza FO, Zerbini PA, Zerbini FM, Fernandes PM (2016) The dsRNA virus Papaya meleira virus and an ssRNA virus are associated with Papaya sticky disease. PLoS One 11(5):e0155240 Saraiva ACM, de Taiva WO, Ravelo Filho FAC, Lima JAA (2006) Transmission by contaminated hands and absence of embryonary seed transmission of Papaya lethal yellowing virus. Fitopatalogia Bras 31:79–83 Saxena S, Hallan V, Singh BP, Sane PV (1998) Leaf curl disease of Carica papaya from India may be caused by a bipartite geminivirus. Plant Dis 82(1):126 Shahid MS, Yoshida S, Khatri-Chhetri GB, Briddon RW, Natsuaki KT (2013) Complete nucleotide sequence of a monopartite begomovirus and associated satellites infecting Carica papaya in Nepal. Virus Genes 46:581–584 Sharma J, Jain RK, Ramiah M, Varma A (2005) Natural spread of Papaya ringspot virus to new areas: occurrence in Coimbatore, Tamil Nadu. Indian Phytopathol 58(2):245–249 Sharma P, Sahu AK, Verma RK, Mishra R, Choudhary DK, Gaur RK (2014) Current status of Potyvirus in India. Arch Phytopathol Plant Prot 47:906–918 Shen W, Tuo D, Yang Y, Yan P, Li X, Zhou P (2014a) First report of Ageratum yellow vein virus associated with a new betasatellite infecting Carica papaya in China. J Plant Pathol 96:603–611 Shen W, Tuo D, Yang Y, Yan P, Li X, Zhou P (2014b) First report of mixed infection of Papaya ringspot virus and Papaya leaf distortion mosaic virus on Carica papaya L. J Plant Pathol 96(4):S4.121 Silva AMR, Kitajima EW, Souza MV, Resende R (1997) Papaya lethal yellowing virus: a possible member of the Tombusvirus genus. Fitopatol Bras 22:529–534 Singh A (2006) Studies of occurrence of papaya viruses in eastern Uttar Pradesh and their possible management approaches. PhD thesis, University of Gorakhpur Singh-Pant P, Pant P, Mukherjee SK, Mazumdar-Leighton S (2012) Spatial and temporal diversity of begomoviral complexes in papayas with leaf curl disease. Arch Virol 157(7):1217–1232 Sinha A, Kumar A, Bhatnagar D, Khan JA (2013) Association of cotton leaf curl Multan virus and its satellite molecules with leaf curl disease of papaya in India. New Dis Rep 27:9 Sit TL, Abouhaidar MG, Holy S (1989) Nucleotide sequence of Papaya mosaic virus RNA. J Gen Virol 70:2325–2331 Somera M, Sarmiento C, Truve E (2015) Overview on Sobemoviruses and a proposal for the creation of the Family Sobemoviridae. Viruses 7:3076–3115 Tapia-Tussell R, Magaña-Alvarez A, Cortes-Velazquez A, Itza-Kuk G, Nexticapan-Garcez A, Quijano-Ramayo A, Martin-Mex R, Perez-Brito D (2015) Seed transmission of Papaya meleira virus in papaya (Carica papaya) cv. Maradol. Plant Pathol 64:272–275 Taya R, Singh J (1995) Survey of Papaya mosaic virus disease in Haryana. Agric Sci Dig 15:191–192 Taya RS, Singh JP (1997) Studies on the transmission of Papaya ring spot virus by Aphis craccivora Koch. Indian J Agric Res 31(4):237–240 Teixeira MGC, de Lima JAA, Sousa AEVA, Fernandes ER (1999) Low incidence levels of Papaya lethal yellowing virus in countries of the Rio Grande do Norte State. Caatinga 12:29–33 Teliz D, Mora G, Nieto D, Gonsalves D, Garcia E, Mathesis I, Oolia C (1991) Papaya ring spot virus in Mexico. Rev Mex Fitopathol 9:64–68 Tennant PF, Fermin GA, Roye ME (2007) Viruses infecting papaya (Carica papaya L.): Etiology, pathogenesis and molecular biology. Plant Viruses 1(2):178–188 Thomas J, Dodman R (1993) The first record of Papaya ringspot virus – type P in Australia. Aust Plant Pathol 22:2–7

Carnegiea gigantea (Saguaro)

455

Tripathi S, Suzuki JY, Ferreira SA, Gonsalves D (2008) Papaya ringspot virus-P: characteristics, pathogenicity, sequence variability and control. Mol Plant Pathol 9(3):269–280 Trujillo EE, Gonsalves D (1967) Tomato spotted wilt in papaya. Phytopathology 57:9 Usharani TR, Laxmi V, Jalali S, Krishnareddy M (2013) Duplex PCR to detect both Papaya ring spot virus and Papaya leaf curl virus simultaneously from naturally infected papaya (Carica papaya L.). Indian J Biotechnol 12:269–272 Verma HN, Prasad V (1985/1986) Virus diseases in pawpaw (papaya). Rev Trop Plant Path 2:311–327 Wan SH, Conover RA (1983) Incidence and distribution of papaya viruses in southern Florida. Plant Dis 67:353–356 Wang HL (1981) Aphid transmission of Papaya ring spot virus in Taiwan. Plant Prot Bull Taiwan 23:229–233 Wang X, Xie Y, Zhou X (2004) Molecular characterization of two distinct Begomoviruses from papaya in China. Virus Genes 29:303–309 Wu CZ, Zhou XP (2006) First report of Malvastrum leaf curl virus infecting papaya. J Plant Pathol 88:342 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Xiao X, Tetsuo M, Luo X, Xiao HG, Tetsuo MK, Luo XH (1997) Investigation and identification of Papaya ringspot virus and Papaya leaf distortion mosaic virus in South China. J South China Agric Univ 18:52–53 Yeh SD, Gonsalves D (1985) Translation of Papaya ring spot virus RNA in vitro detection of a possible polyprotein that is processed for capsid protein cylindrical inclusion protein and amorphous inclusion protein. Virology 143:260–271 Yeh SD, Gonsalves D, Provvidenti R (1984) Comparative studies on host range and serology of Papaya ringspot virus and watermelon mosaic virus 1. Phytopathology 74(9):1081–1085 Yeh SD, Jan FJ, Chiang CH, Doong TJ, Chen MC, Chung PH, Bau HJ (1992) Complete nucleotide sequence and genetic organization of Papaya ringspot virus RNA. J Gen Virol 73(Pt 10):2531–2541 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zhang LB, Zhou GH, Li HP, Zhang SG (2005) Molecular characterization of Papaya leaf curl virus infecting Carica papaya in Guangzhou and its biological test. Sci Agric Sin 38:1805–1810 Zhao H, Jia RZ, Zhang YL, Zhu YJ, Zeng HC, Kong H, McCafferty H, Guo AP, Peng M (2016) Geographical and genetic divergence, Papaya ring spot virus populations within Hainan Province, China. Phytopathology 106:937–944 Zhou X (2013) Advances in understanding begomovirus satellites. Annu Rev Phytopathol 51:357–381

Carnegiea gigantea (Saguaro) Family: Cactaceae

Ornamental

Saguaro cactus virus

(SgCV)

Taxonomic position Genus: Alphacarmovirus

Family: Tombusviridae

Geographical distribution SgCV was first reported in plants of Carnegiea gigantea from Arizona, USA, by Milbrath and Nelson (1972). The virus spreads in cultivated cacti in Germany (Samyn and Welvaert 1978). Symptoms and host(s) The virus-infected saguaro plants show symptomless infections. Transmission The virus is readily transmitted by mechanical sap-inoculation. The virus spreads through vegetative propagation.

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456

Carthamus tinctorius (Safflower)

Virion properties and genome The virions are isometric 30 nm in diameter and have T = 3 icosahedral symmetry. The genome consists of a single molecule of linear positive-sense ssRNA of 3879 nt (U72332 = NC_001780) (Weng and Xiong 1997).

References Milbrath GM, Nelson MR (1972) Isolation and characterization of a virus from saguaro cactus. Phytopathology 62:739–742 Samyn G, Welvaert W (1978) About an isometric virus on a cultivated cactus: Chamaecereus sylvestrii ‘aureus’. Phytopathol Z 91:276–279 Weng Z, Xiong Z (1997) Genome organization and gene expression of Saguaro cactus carmovirus. J Gen Virol 78:525–534

Carthamus tinctorius (Safflower) Family: Asteraceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Oil seed

(AMV)

Family: Bromoviridae

AMV infection in plants of Carthamus tinctorius was reported from India and Serbia (Mali 1998; Milosevic et al. 2015). The virus-infected safflower plants exhibit dark streaks which occur on diseased roots and stems and eventually encircle the stem. Mosaic symptoms may occur on leaves. The virus is transmitted by a large number of aphid species in a non-persistent manner, and also by mechanical sapinoculation. For more details of AMV, refer to Medicago sativa.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Carthamus tinctorius was distrusted wherever the crop is grown (Klisiewicz 1962; Ravinder et al. 1990a, b; Fletcher 2001). The virus-infected safflower plants exhibit symptoms of light and dark green mosaic pattern which occurs on diseased upper leaves and bracts. Some leaves may become blistered and distorted. The mosaic pattern is less distinct on lower leaves and becomes a scattering of light and dark green flecks. Diseased plants may be stunted but usually develop to maturity. The virus is transmitted from diseased to healthy plants early in the growing season by the green peach aphids Myzus persicae and Uroleucon compositae in a non-persistent manner (Ravinder et al. 1990a, b). The virus is also mechanically sap-transmissible. For more details of CMV, refer to Cucumis sativus.

Carthamus tinctorius (Safflower)

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

457

(TMV)

Family: Virgaviridae

TMV infection in plants of Carthamus tinctorius was reported from Morocco (Lockhart and Goethals 1977). The virus-infected safflower plants exhibit symptoms of blotchy light and dark green mosaic patterns which occur on all subsequent leaves until flowering. The virus is transmitted by means not involving a vector. The virus is transmissible by mechanical sap-inoculation. The virus is transmissible by grafting and also through contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Carthamus tinctorius was reported from India and Australia (Chander Rao et al. 2003; Sharman et al. 2015). The virus-infected safflower plants exhibit veinal and leaf necrosis, necrotic streaks on the stem, and necrosis of terminal buds followed by death of the plants. The virus is transmitted by thrips vectors. The virus present in/on pollen, and enters into the host through the injuries caused by thrips while feeding. The virus is also mechanically sap trasmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Carthamus tinctorius was reported from the United States (Klisiewicz 1981). The virus-infected safflower plants exhibit symptoms of stunting, and both leaves and seed heads are reduced in size. Leaves and bracts of seed heads have small dark green and pale green areas, pale green vein-banding, distortion, and bronzing. Seed ovules rot, resulting in a reduction in seed yield. The virus is transmitted by an aphid vector Myzus persicae in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TuMV, refer to Brassica rapa.

References Chander Rao SC, Rao RDVJ, Kumar VN, Raman DS, Raoof MA, Prasad RD (2003) First report of Tobacco streak virus infecting safflower (Carthamus tinctorius) in Maharashtra, India. Plant Dis 87:1396 Fletcher D (2001) New hosts of Alfalfa mosaic virus, Cucumber mosaic virus, Potato virus Y, Soybean dwarf virus, and Tomato spotted wilt virus in New Zealand. N Z J Crop Hortic Sci 29(3):213–217 Klisiewicz JM (1962) Cucumber mosaic virus on safflower. Plant Dis Reptr 46:849–851 Klisiewicz JM (1981) Isolation of Turnip mosaic virus from safflower. Phytopathology 71:886 Lockhart BEL, Goethals M (1977) Natural infection of safflower by a Tobamovirus. Plant Dis Reptr 61:1010–1012

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458

Carya illinoinensis (Pecan)

Mali VR (1998) An appraisal of Alfalfa mosaic alfamovirus (AMV) and AMV diseases in Marathwada (India). Indian J Virol 14:93–104 Milosevic D, Marjanovic-Jeromela A, Jovicic D, Ignjatov M, Nikolic Z, Terzic S, Stankovic I (2015) First report of Alfalfa mosaic virus on safflower in Serbia. Plant Dis 99:896 Ravinder T, Govinda Rao N, Sastry KS (1990a) Relationship of mosaic disease of safflower with is aphid vector Uroleucon compositae. J Insect Sci 3:177–179 Ravinder T, Govinda Rao N, Sastry KS, Sikinder Ali, Ranga Rao V (1990b) Studies on a mosaic disease of safflower in India. In: Proceedings of II international safflower conference held at Hyderabad, pp 285–298 Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207

Carya illinoinensis (Pecan) Family: Juglandaceae

Cherry leaf roll virus Taxonomic position Genus: Nepovirus

Trees/Shrubs

(CLRV)

Family: Secoviridae

CLRV infection in plants of Carya illinoinensis was reported from Syria (Al-Chaabi and Ismaeil 2009). The virus-infected pecan plants do not exhibit obvious symptoms, and only leaf shedding is reported. The virus is transmitted by nematode vectors in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and also by grafting. For more details of CLRV, refer to Prunus avium.

Pecan mosaic-associated virus Taxonomic position Genus: Potyvirus

(PMaV)

Family: Potyviridae

Geographical distribution PMaV infection in plants of Carya illinoensis was reported from China (Su et al. 2016). Symptoms and host(s) The virus-infected pecan plants show leaf mosaic symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long and 11–13 nm wide. The genome contains single molecule of linear, positive-sense single-stranded RNA of 9310 nt in length (KT633868 = NC_030293) (Wylie et al. 2017).

Cassia spp.

459

References Al-Chaabi S, Ismaeil F (2009) First report of the cherry strain of Cherry leaf roll virus on walnut and pecan trees in Syria. J Plant Pathol 91:502 Su X, Fu S, Qian Y, Zhang L, Xu Y, Zhou X (2016) Discovery and small RNA profile of pecan mosaic-associated virus, a novel potyvirus of pecan trees. Sci Rep 26:26741 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Cassia spp. Synonyms Senna spp. Family: Fabaceae

Medicinal

African cassava mosaic virus Taxonomic position Genus: Begomovirus

(ACMV)

Family: Geminiviridae

ACMV infection in plants of Senna occidentalis was reported from Nigeria (Ogbe et al. 2006; Alabi et al. 2008). The virus-infected senna plants exhibit mosaic symptoms. The virus is transmitted by a whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ACMV, refer to Manihot esculenta.

Bean common mosaic necrosis virus Taxonomic position Genus: Potyvirus

(BCMNV)

Family: Potyviridae

BCMNV infection in plants of Senna spp. was reported from Uganda by Sengooba et al. (1997). The virus-infected senna plants exhibit chlorotic spots, chlorotic streaks, and mild mosaic symptoms. The virus is transmitted by an aphid vector Aphis fabae in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BCMNV, refer to Phaseolus vulgaris.

Cassia severe mosaic virus

(CSMV)

Taxonomic position CSMV is a tentative member of the genus Potyvirus in the family Potyviridae Geographic distribution CSMV was reported from the Yemen Republic and Ethiopia (Walkey et al. 1994).

C

460

Cassia spp.

Symptoms and host range The virus-infected cassia plants showed severe mosaic symptoms; cassia was the only natural host identified, with a limited experimental range of a few legumes, plus Nicotiana clevelandii and N. benthamiana. Transmission The virus was transmitted by the aphid Myzus persicae in a non-persistent manner, and also by mechanical inoculation with sap. Virion properties and genome Flexuous rod-shaped particles of mean length of 830 nm were observed, and pinwheels and scrolls typical of potyviruses were observed in thin sections. The viral coat protein of c.34.5 kDa reacted with a monoclonal antibody with broad spectrum reactivity against potyviruses, and with a virus-specific polyclonal antiserum, but not against polyclonal antisera specific for eight other potyviruses including bean common mosaic virus, bean yellow mosaic virus, clover yellow vein virus, cowpea aphid-borne mosaic virus, peanut mottle virus, and several potyviruses not normally infecting leguminous hosts (Walkey et al. 1994). The genome is presumed to be single-stranded RNA of 9.5–10 kb, but no sequence information is available.

Cassia yellow blotch virus Taxonomic position Genus: Bromovirus

(CYBV)

Family: Bromoviridae

Geographical distribution CYBV infection was first reported in plants of Cassia pleurocarpa from Mitchell, Queensland, by Dale et al. (1984). The virus spreads in Australia (Lenne 1990). Symptoms and host(s) The virus-infected cassia plants exhibit yellow leaf blotches and leaf deformations. Transmission The virus is transmitted by beetle vectors. The virus is also transmissible by mechanical sap-inoculation. Under experimental conditions, CYBV infects Arabidopsis thaliana causing systemic symptoms. Virion properties and genome The virions are non-enveloped isometric particles about 26 nm in diameter with T = 3 icosahedral symmetry, composed of 180 capsid proteins: 12 pentamers and 20 hexamers. The genome is segmented, tripartite linear positive-sense ssRNA composed of RNA1 (3176 nt, AB194806 = NC_006999), RNA2 (2721 nt, AB194807 = NC_007000), and RNA3 (2091 nt, AB194808 = NC_007001). Each genomic segment has a 30 tRNA-like structure and a 50 cap (Iwahashi et al. 2005; Kao and Adkins 2011; Scott 2011).

Cassia yellow mosaic-associated virus Taxonomic position The virus is a tentative member of the genus Tymovirus and family Tymoviridae.

Cassia spp.

461

Geographical distribution The virus-infected Cassia hoffmannseggii plants were reported from Brazil (Nicolini et al. 2012). Symptoms and host(s) The virus-infected cassia plants exhibit severe mosaic symptoms. Transmission The virus is transmitted by beetle vectors in a semi-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are isometric of c.30 nm diameter, including both ‘full’ and ‘empty’ particles. A partial sequence of 1653 nt is available (JN545837).

East African cassava mosaic virus Taxonomic position Genus: Begomovirus

(EACMV)

Family: Geminiviridae

EACMV infection in plants of Senna occidentalis was reported from Nigeria (Ogbe et al. 2006; Alabi et al. 2008). The virus-infected senna plants exhibit mosaic symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of EACMV, refer to Manihot esculenta.

Senna chlorotic stunt virus

(SeCSV)

Taxonomic position SeCSV is a tentative member of the family Closteroviridae. Geographical distribution SeCSV infection in plants of Senna pallida was reported from Venezuela (Marys et al. 2000). Symptoms and host(s) The virus-infected senna plants exhibit symptoms of stunting, mosaic, vein yellowing, and leaf malformation. Transmission The virus is mechanically sap-transmissible to Senna pallida, Cassia obovata, and Cassia emarginata L. Virion properties and genome The virions are flexuous thread-like particles c. 650–700 nm in length. The genome is linear positivesense single-stranded RNA; dsRNAs of c.12.3 kb, 10 kb, and 5.4 kb were detected from infected plants. The coat protein was shown to be of 26–28 kDa, and to be serologically related to that of the crinivirus lettuce infectious yellows virus (Marys et al. 2000).

C

462

Senna leaf curl virus Taxonomic position Genus: Begomovirus

Cassia spp.

(SenLCuV)

Family: Geminiviridae

Geographical distribution SenLCuV infection in plants of Senna occidentalis was reported from India (Kumar et al. 2016). Symptoms and host(s) The virus-infected senna plants exhibit leaf curl symptoms. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative nonpropagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2742 nt (KU852742 = NC_030748) (Briddon 2001; Brown et al. 2015; Kumar et al. 2016; Zerbini et al. 2017).

Senna mosaic virus

(SenMV)

Taxonomic position SenMV is a tentative member of the genus Potexvirus and family Alphaflexiviridae Geographical distribution SenMV infection in plants of Senna occidentalis and S. hirsuta was reported from Brazil and Nigeria (Owolabi and Proll 2001; Rezende et al. 2017). Symptoms and host(s) The virus-infected senna plants exhibit mosaic and severe leaf malformation symptoms. Transmission The virus was transmissible to S. occidentalis by mechanical sap-inoculation, and trimming scissors, but not by the seeds. Virion properties and genome The complete genome sequence of SenMV comprises 6775 nt excluding the poly(A) tail with five open reading frames coding for RNA-dependent RNA polymerase (RdRp), the triple gene block (TGB 1, 2, and 3) proteins, and coat protein (CP) (Rezende et al. 2017).

Senna virus X Synonyms Cassia mild mosaic virus

Cassia spp.

463

Taxonomic position The virus is a tentative Unassigned speciesof the order Tymovirales Geographical distribution The virus infection in plants of Senna macranthera was reported from Brazil (Beserra et al. 2011). Senna virus X was previously described as Cassia mild mosaic virus from Brazil (Lin et al. 1979, 1980). Symptoms and host(s) The virus-infected senna plants exhibit symptoms of chlorotic spots in the young leaves and mild mosaic in the old leaves. Transmission The virus is transmissible by mechanical sap-inoculation to less than three families. The virus is not transmissible by contact between plants. The virus is not transmissible through the seed and pollen. Virion properties and genome The virions are flexuous filaments, non-enveloped, with a clear modal length of 640 nm and 15 nm wide. The genome is single-stranded RNA.

Senna virus Y Synonyms Cassia yellow spot virus Taxonomic position The virus is a tentative member of the genus Potyvirus and family Potyviridae. Geographical distribution The virus infection in Senna macranthera plants was reported from Brazil (Beserra et al. 2011). Senna virus Y was previously described as Cassia yellow spot virus from Brazil (Paguio and Kitajima 1981; Souto and Kitajima 1991). Symptoms and host(s) The virus-infected senna plants exhibit symptoms of chlorotic spots in the young leaves and mild mosaic in the old leaves. Transmission The virus is transmitted by aphid vectors, Myzus persicae and Uroleucon ambrosiae, in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation. The virus is transmissible through the seed of Cassia occidentalis (Souto 1990). Virion properties and genome The virions are flexuous filaments and non-enveloped, with a clear modal length of 700 nm. The genome is a single molecule of positive-sense single-stranded RNA, probably of 9.5–10 kb.

C

464

Cassia spp.

Southern bean mosaic virus Taxonomic position Genus: Sobemovirus

(SBMV)

Family: Solemoviridae

SBMV infection in plants of Cassia spp. was reported from Lome, Togo (West Africa), by Gumedzoe (1993). The virus was also reported from India (Singh and Singh 1973). The virus-infected cassia plants exhibited mosaic mottling symptoms. The virus is transmitted by beetle vectors such as Cerotoma trifurcata and Epilachna varivestis in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of SBMV, refer to Phaseolus vulgaris.

Soybean mosaic virus Taxonomic position Genus: Potyvirus

(SMV)

Family: Potyviridae

SMV-infected Senna occidentalis (Syn. Cassia occcidentalis) plants were reported from Brazil and India (Singh and Gupta 1996; Almeida et al. 2002). The virus-infected senna plants exhibit mosaic and blistering symptoms. The virus is transmitted by aphid vectors, Aphis gossypii and Taxoptera citricida, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SMV, refer to Glycine max.

Tomato leaf curl Patna virus Taxonomic position Genus: Begomovirus

(ToLCPatV)

Family: Geminiviridae

ToLCPatV infection in plants of Cassia tora was reported from India (Sohrab 2016). The virus-infected cassia plants exhibit leaf curling symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of ToLCPatV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Cassia spp. was reported from Lome, Togo (West Africa), by Gumedzoe (1993). The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation, and by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Cassia spp.

465

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Senna obtusifolia was reported from Georgia (USA) (Mullis et al. 2009). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Alabi OJ, Ogbe FO, Bandyopadhyay R, Lava Kumar P, Dixon AGO, Hughes J’A, Naidu RA (2008) Alternate hosts of African cassava mosaic virus and East African cassava mosaic Cameroon virus in Nigeria. Arch Virol 153:1743–1747 Almeida AMR, Sakai J, Souto ER, Kitajima EW, Fukuji TS, Hanada K (2002) Mosaic in Senna occidentalis in southern Brazil induced by a new strain of Soybean mosaic virus. Fitopathol Bras 27:151–156 Beserra JEA Jr, de Carvalho MG, Barguil BM, Zerbini FM (2011) Partial genome sequence of a potyvirus and of a virus in the order Tymovirales found in Senna macranthera in Brazil. Trop Plant Pathol 36:116–120 Briddon RW (2001) Begomovirus. Geminiviridae. The Springer Index of Viruses. Springer, New York, pp 340–348, https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Dale JL, Gibbs AJ, Behncken GM (1984) Cassia yellow blotch virus, a new bromovirus from an Australian native legume Cassia pleurocarpa. J Gen Virol 65:281–288 Gumedzoe MY (1993) Major virus diseases of medicinal plants in west Africa. Acta Hortic 331:307–310 Iwahashi F, Fujisaki K, Kaido M, Okuno T, Mise K (2005) Synthesis of infectious in vitro transcripts from Cassia yellow blotch bromovirus cDNA clones and a reassortment analysis with other bromoviruses in protoplasts. Arch Virol 150:1301–1314 Kao CC, Adkins S (2011) Bromovirus. Bromoviridae. In: The Springer index of Viruses. Springer, New York, pp 173–177. https://doi.org/10.1007/978-0-387-95919-1_25 Kumar J, Alok A, Kumar J, Tuli R (2016) Senna leaf curl virus: a novel Begomovirus identified in Senna occidentalis. Arch Virol 161:2609–2612 Lenne JM (1990) Diseases of Cassia species – a review. Trop Grasslands 24:311–324 Lin MT, Kitajima EW, Cupertino FP, Costa CL (1979) Properties of a possible carlavirus from a cerrado native plant Cassia sylvestris. Plant Dis Reptr 6:501–505 Lin MT, Kitajima EW, Costa CL (1980) Association of Cassia mild mosaic virus with dieback of Cassia macranthera in Central Brazil. Plant Dis Reptr 64:587–589 Marys EE, Liu H-Y, Carballo O, Romano M, Uzcategui R, Colmenares M, Izaguirre-Mayoral ML (2000) Partial characterisation of a new virus in brusca from the Venezuelan lowlands. Ann Appl Biol 136:125–130 Mullis S, Bertrand PF, Brown SL, Csinos A, Diaz-Perez JC, Gitaitis R, Hickman LL, Johnson A, LaHue SS, Martinez N, McPherson RM, Nischwitz C, Reay-Jones F, Riley D, Sherwood J, Stevenson K, Wells L (2009) Tospoviruses in Solanaceae and other crops in the coastal plain of Georgia. University of Georgia, College of Agricultural and Environmental Sciences, Bulletin 1354, pp 51 Nicolini C, Pio-Ribeiro G, Andrade GP, Melo FL, Oliveira VC, Guimarães FC, Resende RO, Kitajima EW, Rezende JA, Nagata T (2012) A distinct tymovirus infecting Cassia hoffmannseggii in Brazil. Virus Genes 45:190–194 Ogbe FO, Dixon AGO, Hughes JA, Alabi OJ, Okechukw R (2006) Status of cassava begomo viruses and their new natural hosts in Nigeria. Plant Dis 90:548–553 Owolabi AT, Proll E (2001) A mosaic disease of Senna hirsuta induced by a potyvirus in Nigeria. Acta Virol 45:73–79 Paguio DR, Kitajima EW (1981) Isolamento de um potyvirus de Cassia hoffmannsegii Mart. Fitopatol Bras 6:187–191 Rezende JAM, Camelo-Garcia VM, Andrade SCS, Buriolla JE, Kitajima EW, Duarte LML (2017) Biological and molecular characterization of a putative new potexvirus infecting Senna occidentalis. Arch Virol 162:529–533 Scott SW (2011) Bromoviridae and allies. In: Harper D (ed) Encyclopedia of Life Sciences (ELS). Wiley, Chichester. https://doi.org/10.1002/9780470015902

C

466

Catalpa bignonioides (Indian bean tree)

Sengooba TN, Spence NJ, Walkey DGA, Allen DJ, Femi Lana A (1997) The occurrence of Bean common mosaic necrosis virus in wild and forage legumes in Uganda. Plant Pathol 46:95–103 Singh BR, Gupta SP (1996) Natural occurrence of Soybean mosaic virus on coffee senna. Indian J Virol 12:71–72 Singh R, Singh R (1973) Natural infection of sickle senna (Cassia tora L.) and cowpea (Vigna sinensis savi) plants by some new strains of Southern bean mosaic virus. Port Acta Biol 13:87–98 Sohrab SS (2016) First report of Tomato leaf curl Patna virus infecting Cassia tora in India. Plant Dis 100:1798 Souto ER (1990) Cassia yellow spot virus. Inst Clienc Biol Univ de Brasilia, p 93 Souto ER, Kitajima EW (1991) Propriedades biológicas de um potyvírus isolado de “Lava Pratos” (Cassia hoffmannseggii) procedente de Pernambuco. Fitopatol Bras 16:256–259 Walkey DGA, Spence NJ, Clay CM, Miller A (1994) A Potyvirus isolated from Senna occidentalis. Plant Pathology 43:767–773 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

Catalpa bignonioides (Indian bean tree) Family: Bignoniaceae

Broad bean wilt virus Taxonomic position Genus: Fabavirus

Trees/Shrubs

(BBWV)

Family: Secoviridae

BBWV (but without specifying either of the currently recognized species BBWV-1 or BBWV-2) infection in plants of Catalpa bignonioides was reported from the Eastern Europe (Schmelzer 1970). The virus-infected Indian bean tree plants exhibit chlorotic leaf spotting symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BBWV, refer to Vicia faba.

References Schmelzer K (1970) Investigations on the viruses of ornamental and wild woody plants. 7th report: further reports on Buddleia, Viburnum, Caryopteris and Philadelphus, as well as virus diseases of Leycesteria, Chionanthus, Ribes, Hydrangea, Syringa, Spiraea and Catalpa. Phytopathol Z 67:283–326

Catharanthus spp. (Periwinkle) Synonyms Vinca spp. Family: Apocynaceae

Ornamental

Catharanthus spp. (Periwinkle)

467

Arabis mosaic virus

(ArMV)

Taxonomic position Genus: Nepovirus

Family: Secoviridae

ArMV infection in plants of Vinca spp. was reported from Ohio, USA (Fisher 2013). The virus-infected periwinkle plants exhibit mottle and marginal/interveinal chlorosis of leaves. The virus is transmitted by the dagger nematode vector, Xiphinema diversicaudatum, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Broad bean wilt virus 2 Taxonomic position Genus: Fabavirus

(BBWV-2)

Family: Secoviridae

BBWV-2 infection in plants of Vinca rosea was reported from Australia (Shukla et al. 1980). The virusinfected periwinkle plants do not exhibit any symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BBWV-2, refer to Vicia faba.

Catharanthus mosaic virus Taxonomic position Genus: Potyvirus

(CatMV)

Family: Potyviridae

Geographical distribution CatMV infection in plants of Catharanthus roseus was reported from Brazil and Saudi Arabia (Conceicao et al. 2011; Maciel et al. 2011; Elbeshehy et al. 2017). Symptoms and host(s) The virus-infected periwinkle plants show mosaic, foliar malformation, and flower variegation symptoms. Transmission The virus is transmitted by aphid vectors, Aphis gossypii and Myzus nicotianae, in a non-persistent manner. The virus is also mechanically sap-transmissible to Nicotiana benthamiana, Chenopodium amaranticolor, and C. quinoa. Virion properties and genome The virions are non-enveloped, flexuous filaments, 700–900 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA and consists of 9575 nt (KP343681 = NC_027210) (Revers and Garcia 2015; Wylie et al. 2017).

C

468

Catharanthus spp. (Periwinkle)

Catharanthus yellow mosaic virus Taxonomic position Genus: Begomovirus

(CatYMV)

Family: Geminiviridae

Geographical distribution CatYMV infection in plants of Catharanthus roseus was reported from Pakistan and Saudi Arabia (Ilyas et al. 2013; Elbeshehy et al. 2017). Symptoms and host(s) The virus-infected periwinkle plants exhibit irregular yellow mosaic symptoms with severe curling and distortion. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2752 nt (LK028570 = NC_025964) (Briddon 2001; Ilyas et al. 2013; Brown et al. 2015; Zerbini et al. 2017).

Chrysanthemum stunt viroid Taxonomic position Genus: Pospiviroid

(CSVd)

Family: Pospiviroidae

CSVd infection in plants of Vinca major was reported from Canada (Nie et al. 2005). The viroidinfected periwinkle plants do not exhibit any symptoms. The viroid is mechanically sap-transmissible. For more details of CSVd, refer to Chrysanthemum spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Catharanthus spp. was reported from Saudi Arabia, the USA, Brazil, Australia, Korea, Malaysia, and elsewhere (Shukla et al. 1980; Smith and Davis 1986; Giorgadze et al. 1988; Espinha and Gaspar 1997; Samad et al. 2008; Fisher 2012; Mazidah et al. 2012; Choi et al. 2014; Elbeshehy et al. 2017). The virus-infected periwinkle plants show bright chlorotic spots on the upper leaves in early summer, progressing to green mosaic, deformed leaves, and stunted growth; flowers may be malformed with faded petals. The virus is transmitted by an aphid vector Myzus persicae in a nonpersistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Catharanthus spp. (Periwinkle)

469

Groundnut bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(GBNV)

Family: Tospoviridae

GBNV infection in plants of Catharanthus roseus was reported from India (Daimei et al. 2017; Basavaraj et al. 2017). The infected periwinkle plants exhibited stunted growth, bud necrosis, necrotic spots, and veinal necrosis on leaves, leaf distortion, stem necrosis, and even death of the plant in severe cases. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of GBNV, refer to Arachis hypogaea.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Catharanthus roseus was reported from Italy and the USA (Ruter and Gitaitis 1993a). The virus-infected periwinkle plants exhibit symptoms such as black foliar lesions, distortion of young growth, stunting, and wilting. These symptoms can vary with the stage of growth of the host and with a variety of cultural conditions. The virus is transmitted by a thrips vector Frankliniella occidentalis in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Iresine viroid 1

(IrVd-1)

Taxonomic position Genus: Pospiviroid

Family: Pospiviroidae

IrVd-1 infection in plants of Vinca major was reported from Canada (Nie et al. 2005). The viroidinfected periwinkle plants do not exhibit any obvious symptoms. The viroid is mechanically saptransmissible. For more details of IrVd-1, refer to Iresine spp.

Lettuce chlorosis virus Taxonomic position Genus: Crinivirus

(LCV)

Family: Closteroviridae

LCV infection in plants of Catharanthus roseus was reported from China (Tian et al. 2018). The virusinfected periwinkle plants exhibit symptoms of interveinal yellowing, leaf curling, and plant dwarfing. The virus is transmitted by whitefly vectors in a semi-persistent manner. The virus is not mechanically sap-transmissible. For more details of LCV, refer to Lactuca sativa.

C

470

Lettuce mosaic virus Taxonomic position Genus: Potyvirus

Catharanthus spp. (Periwinkle)

(LMV)

Family: Potyviridae

LMV infection in plants of Catharanthus roseus was reported from France and Saudi Arabia (SvanellaDumas et al. 2014; Elbeshehy et al. 2017). The virus-infected periwinkle plants exhibit symptoms of deformation and dark and light green sectoring of the leaves along the major veins and a modification of flower morphology, with more slender and no longer overlapping petals. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of LMV, refer to Lactuca sativa.

Potato yellow dwarf nucleorhabdovirus Taxonomic position Genus: Nucleorhabdovirus

(PYDV)

Family: Rhabdoviridae

PYDV infection in periwinkle (Vinca rosea ) plants was reported from California (Falk et al. 1981). The virus-infected periwinkle plants exhibit symptoms of chlorosis, with a mosaic pattern in older leaves and twisting and necrosis of very young leaves. The virus is transmitted by leafhopper vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of PYDV, refer to Solanum tuberosum.

Tomato chlorotic dwarf viroid Taxonomic position Genus: Pospiviroid

(TCDVd)

Family: Pospiviroidae

TCDVd infection in plants of Vinca minor was reported from the USA, Belgium, and Canada (Singh and Dilworth 2009; Van Bogaert et al. 2017). The viroid-infected periwinkle plants do not exhibit any symptoms. The viroid is mechanically sap-transmissible. For more details of TCDVd, refer to Solanum lycopersicum.

Tomato chlorotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(TCSV)

Family: Tospoviridae

TCSV infection in plants of annual vinca (Catharanthus roseus) was reported from the USA (Warfield et al. 2015). The virus-infected periwinkle plants exhibit symptoms of dark brown lesions on the stems, chlorotic leaves with necrotic line patterns, spots, ringspots, and black petioles. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of TCSV, refer to Solanum lycopersicum.

Catharanthus spp. (Periwinkle)

471

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Catharanthus roseus was reported from Saudi Arabia, Portugal, Greece, Canada, and the USA (Paliwal 1974; Ruter and Gitaitis 1993b; Louro 1996; Chatzivassiliou et al. 2000; Elbeshehy et al. 2017). The virus-infected periwinkle plants show black spots, systemic mosaic, leaf deformation, and browning of larger leaves at the lower part of the plant. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Vinca leaf curl virus

(VinLCV)

Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution VinLCV infection in plants of Vinca rosea was reported from India (Nehra et al. unpublished; KR612272). Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consist of a single molecule of circular single-stranded DNA (DNA-A) of 2776 nt (KR612272 = NC_028124) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV was identified from India in plants of Catharanthus roseus showing severe chlorosis, crinkling, and curling of leaves (Srivastava et al. 2012, 2013). The virus is readily transmitted mechanically and by multiple species of aphids (e.g., Aphis gossypii, A. nerii, A. craccivora, Brevicoryne brassicae, Myzus persicae, and Macrosiphoniella sanborni) in a non-persistent manner. For more details of WMV, refer to Citrullus lanatus.

References Basavaraj YB, Kumar A, Holkar SK, Jain RK, Mandal B (2017) First report of Groundnut bud necrosis virus infecting periwinkle (Catharanthus roseus) in India. Plant Dis 101:1559 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of plant Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3–540-31042-8_55

C

472

Catharanthus spp. (Periwinkle)

Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Chatzivassiliou E, Livieratos I, Jenser G, Katis N (2000) Ornamental plants and thrips populations associated with Tomato spotted wilt virus in Greece. Phytoparasitica 28:257–264 Choi S-K, Cho I-S, Choi G-S, Yoon J-Y (2014) First report of Cucumber mosaic virus in Catharanthus roseus in Korea. Plant Dis 98:1283 Conceicao MS, Da Silva RF, Reis MS, Jadão AS, Rosa DD, Giampan JS, Kitajima EW, Rezende JAN, Camargo LEA (2011) Characterization of a new potyvirus causing mosaic and flower variegation in Catharanthus roseus in Brazil. Sci Agric 68:687–690 Daimei G, Pukhrambam PD, Saurav GK, Raina HS, Rajagopal R (2017) First report of the natural occurrence of Groundnut bud necrosis virus on Catharanthus roseus in India. Plant Dis 101:1333 Elbeshehy KF, Al-Zahrani HSM, Aldhebiani AY, Elbeaino T (2017) Viruses infecting periwinkle (Catharanthus roseus L.) in western Saudi Arabia. Phytopathol Mediterr 56:479–485 Espinha LM, Gaspar JO (1997) Partial characterization of Cucumber mosaic virus isolated from Catharanthus roseus. Fitopatol Bras 22:209–212 Falk BW, Weathers LG, Greer FC (1981) Identification of Potato yellow dwarf virus occurring naturally in California. Plant Dis 65:81–83 Fisher JR (2012) Identification of three distinct classes of satellite RNAs associated with two Cucumber mosaic virus serotypes from the ornamental ground cover Vinca minor. Plant Health Prog. https://doi.org/10.1094/PHP-2012-041201-RS. Online Fisher JR (2013) First report of Arabis mosaic virus infecting Vinca minor in Ohio. Plant Health Prog. https://doi.org/ 10.1094/PHP-2013-0901-02-BR. Online Giorgadze RD, Nikolaishvili AA, Dzimistarishvili NB (1988) Cucumber mosaic virus on pink Catharanthus. Subtropicheskie Kult 1:132–135 Ilyas M, Nawaz K, Shafiq M, Haider MS, Shahid AA (2013) Complete nucleotide sequences of two begomoviruses infecting Madagascar periwinkle (Catharanthus roseus) from Pakistan. Arch Virol 158:505–510 Louro D (1996) Detection and identification of Tomato spotted wilt virus and Impatiens necrotic spot virus in Portugal. Acta Hortic 431:99–105 Maciel SC, Silva RF, Reis MS, Jadao AS, Rosa DD, Giampan JS, Kitajima EW, Rezende JAM, Camargo LEA (2011) Characterization of a new potyvirus causing mosaic and flower variegation in Catharanthus roseus in Brazil. Sci Agric 68:687–690 Mazidah M, Yusoff K, Habibuddin H, Tan YH, Lau WH (2012) Characterization of Cucumber mosaic virus (CMV) causing mosaic symptom on Catharanthus roseus (L.) G. Don in Malaysia. Pertanika J Trop Agric Sci 35:41–53 Nie X, Singh RP, Bostan H (2005) Molecular cloning, secondary structure, and phylogeny of three pospiviroids from ornamental plants. Can J Plant Pathol 27:592–602 Paliwal YC (1974) Some properties and thrip transmission of Tomato spotted wilt virus in Canada. Can J Bot 52:1177–1182 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Ruter JM, Gitaitis RD (1993a) Impatiens necrotic spot virus in woody landscape plants in Georgia. Plant Dis 77:318 Ruter JM, Gitaitis RD (1993b) First report of Tomato spotted wilt virus on bedding plants in Georgia. Plant Dis 77:101 Samad A, Ajayakumar PV, Gupta MK, Shukla AK, Darokar MP, Somkuwar B, Alam M (2008) Natural infection of periwinkle (Catharanthus roseus) with Cucumber mosaic virus, subgroup IB. Australas Plant Dis Notes 3:30–34 Shukla DD, Teakle DS, Gough KH (1980) Periwinkle, a latent host for broad bean wilt and cucumber mosaic viruses in Australia. Plant Dis 64:802–803 Singh RP, Dilworth AD (2009) Tomato chlorotic dwarf viroid in the ornamental plant Vinca minor and its transmission through tomato seed. Eur J Plant Pathol 123(1):111–116 Smith DB, Davis RF (1986) Identification of Cucumber mosaic virus from Vinca minor in New Jersey. Plant Dis 70:1056–1059 Srivastava D, Panday N, Tiwari AK, Shukla K (2012) Identification of a potyvirus associated with mosaic disease of Catharanthus roseus and its histopathogical effects. Med Plants 4:23–27 Srivastava D, Pandey N, Tiwari AK, Shukla K (2013) Identification of a Watermelon mosaic virus associated with mosaic disease of Catharanthus roseus in North-Eastern Uttar Pradesh and its histopathological effects. Indian J Virol 24:148 Svanella-Dumas L, Verdin E, Faure C, German-Retana S, Gognalons P, Danet JL, Marais A, Candresse T (2014) Adaptation of Lettuce mosaic virus to Catharanthus roseus involves mutations in the central domain of the VPg. Mol Plant-Microbe Interact 27:491–497 Tian X, Tian Y, Yu YQ, Wang XF, Li ZA, Li RH, Cao MJ, Zhou CY (2018) Periwinkle: a new natural host of Lettuce chlorosis virus in China. Plant Dis 102:462 Van Bogaert N, Smagghe G, Maes M, De Backer M, De Jonghe K (2017) Phylogeny of five predominant pospiviroid species in Belgium. Eur J Plant Pathol 149:25–33

Cattleya spp.

473

Warfield CY, Clemens K, Adkins S (2015) First report of Tomato chlorotic spot virus on annual vinca (Catharanthus roseus) in the United States. Plant Dis 99:895 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

C Cattleya spp. Synonyms Laelia spp. Family: Orchidaceae

Ornamental

Cymbidium mosaic virus Taxonomic position Genus: Potexvirus

(CymMV)

Family: Alphaflexiviridae

CymMV infection in plants of Cattleya spp. was reported from Puerto Rico, Ecuador, Florida, Thailand, and Hawaii (Zettler et al. 1990; Hu et al. 1993; Elliott et al. 1996; Khentry et al. 2006). The virusinfected cattleya plants exhibit symptoms of brown to purple streaks or flecks that appear; in severe cases, older leaves drop prematurely leaving the plant with two or three leaves, and the balance of the plant becomes leafless pseudobulbs. The first symptom on the new growth appears as yellow blotched areas that may be oval or elongated streaks. At times the plant may be infected with this virus without noticeable symptoms on any part of the old or new growth. Flowers of Cattleya-type orchids develop brown necrotic streaks or necrosis 5–21 days after they open (Faccioli and Marani 1979). The virus is mechanically sap-transmissible and also by contact between plants. No vector transmission is reported. For more details of CymMV, refer to Cymbidium spp.

Laelia red leafspot virus

(LRLV)

Taxonomic position LRLV is a tentative member of the family Rhabdoviridae. Geographical distribution LRLV infection in plants of laelia was reported from Germany (Peters 1977). Symptoms and host(s) Plants of Laelia purpurata and Laeliocattleya “Isenteana” show dark red spots and ringspots on maturing leaves, with a diameter of 1–5 mm, while fully mature leaves show sunken areas of similar dimensions but without red coloration.

474

Cayaponia tibiricae

Virion properties and genome The virions are bacilliform, 280–315 nm in length and 80 nm wide. The genome is presumed to be single-stranded negative-sense RNA of >12 kb.

Odontoglossum ringspot virus Taxonomic position Genus: Tobamovirus

(ORSV)

Family: Virgaviridae

ORSV infection in plants of Cattleya spp. was reported from Puerto Rico, Ecuador, Thailand, and the United States (Zettler et al. 1990; Hu et al. 1993; Elliott et al. 1996; Khentry et al. 2006). The virusinfected cattleya plants exhibit symptoms of flower color-breaking, chlorotic streaking, mosaic, or necrosis. The virus is transmitted by means not involving a vector. The virus is transmissible by mechanical sap-inoculation, and also through contact between plants. For more details of ORSV, refer to Odontoglossum grande.

References Elliott MS, Zettler FW, Zimmerman MT, Barnett OW Jr, Le Grande MD (1996) Problems with interpretation of serological assays in a virus survey of orchid species from Puerto Rico, Ecuador, and Florida. Plant Dis 80:1160–1164 Faccioli G, Marani F (1979) Cymbidium mosaic virus associated with flower necrosis in Cattleya orchids. Phytopath Med 18:21–25 Hu J, Ferreira S, Wang M, Xu MQ (1993) Detection of Cymbidium mosaic virus, Odontoglossum ringspot virus, Tomato spotted wilt virus, and potyviruses infecting orchids in Hawaii. Plant Dis 77:464–468 Khentry Y, Paradornuwat A, Tantiwiwat S, Phansiri S, Thaveechai N (2006) Incidence of Cymbidium mosaic virus and Odontoglossum ringspot virus on in vitro Thai native orchid seedlings and cultivated orchid mericlones. Kasetart J (Nat Sci) 40:49–57 Peters K-R (1977) Orchid viruses, a new rhabdovirus in Laelia red leafspots. J Ultrastruct Res 58:166–171 Zettler FH, Ko NJ, Wisler GC, Eliote MS, Wong SM (1990) Viruses of orchids and their control. Plant Dis 74:621–626

Cayaponia tibiricae Family: Cucurbitaceae

Weed host

Zucchini yellow mosaic virus Taxonomic position Genus: Potyvirus

(ZYMV)

Family: Potyviridae

ZYMV infection in plants of Cayaponia tibiricae was reported from Brazil (Yuki et al. 1999). The virus-infected plants exhibit yellow mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ZYMV, refer to Cucurbita pepo.

Celosia spp. (Cockscomb)

475

References Yuki VA, Rezende JAM, Kitajima EW, Barroso PAV, Kuniyuki H, Groppo GA, Pavan MA (1999) Cayaponia tibiricae: new host of Zucchini yellow mosaic virus in Brazil. Plant Dis 83:486

Celosia spp. (Cockscomb) Family: Amaranthaceae

C Ornamental

Alternanthera mosaic virus Taxonomic position Genus: Potexvirus

(AltMV)

Family: Alphaflexiviridae

AltMV infection in plants of Celosia spp. was reported from the USA (Hammond and Rane unpublished). The virus-infected cockscomb plants show mosaic, leaf distortion, and reddening symptoms. No vector is reported for this virus. The virus is readily sap-transmissible and easily spread by plant contact, vegetative propagation, and contaminated tools and hands. For more details of AltMV, refer to Alternanthera spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Celosia argentea was reported from commercial greenhouses in the USA and India (Provvidenti 1975; Raj et al. 2000). The virus-infected cockscomb plants show mottle or mosaic symptoms. The virus is transmitted by aphid vectors Aphis craccivora and A. gossypii in a nonpersistent manner, and also by mechanical sap-inoculation to a large number of host plants. For more details of CMV, refer to Cucumis sativus.

Iresine viroid 1

(IrVd-1)

Taxonomic position Genus: Pospiviroid

Family: Pospiviroidae

IrVd-1 isolate was reported in pants of Celosia spp. in the Netherlands and Italy (Verhoeven et al. 2010; Sorrentino et al. 2015). The virus-infected cockscomb plants do not exhibit any symptoms. The viroid is transmissible mechanically to C. plumosa. For more details of IrVd-1, refer to Iresine spp.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

476

Centaurea spp.

TRSV was identified in pants of Celosia spp. in the Netherlands (Gera et al. 2011). The virus-infected cockscomb plants exhibit ringspot symptoms. The virus is transmitted by nematode vectors in a nonpersistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in pants of Celosia cristata was reported from Pennsylvania (USA) (Hausbeck et al. 1992). The virus-infected cockscomb plants exhibit mottle mosaic, ringspots, chlorosis, and necrosis symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Gera A, Beckelman E, Katzir P, Tam Y, Spiegel S, Zeidan M (2011) New and emerging viruses in ornamental crops. Acta Hortic 901:105–112 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among greenhouse ornamentals in Pennsylvania. Plant Dis 76:795–800 Provvidenti R (1975) Natural infection of Celosia argentea by Cucumber mosaic virus in a commercial greenhouse. Plant Dis Reptr 59:166–168 Raj SK, Aminuddin, Pandey RM, Nath P (2000) Screening and field evaluation for natural occurrence of Cucumber mosaic virus in Amaranths and Celosia. Indian J Plant Pathol 18:14–18 Sorrentino R, Minutolo M, Alioto D, Torchetti EM, Di Serio F (2015) First report of Iresine viroid 1 in ornamental plants in Italy and of Celosia cristata as a novel natural host. Plant Dis 99:1655 Verhoeven JTHJ, Jansen CCC, Botermans M, Roenhorst JW (2010) First report of Iresine viroid 1 in Celosia plumosa in the Netherlands. Plant Dis 94:920

Centaurea spp. Family: Asteraceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in pants of Centaurea cyanus was reported from China (Tien-Po et al. 1982). The virus-infected centaurea plants exhibit severe mosaic and leaf deformation symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Centranthus ruber (Red valerian)

477

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in pants of Centaurea spp. was reported by Parrella et al. (2003). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Parrella G, Gognalons P, Gebre-Selassi K, Vovlas C, Marchoux G (2003) An update of the host range of Tomato spotted wilt virus. J Plant Pathol 85:227–264 Tien-Po, Rao ALN, Hatta T (1982) Cucumber mosaic virus from cornflower in China. Plant Dis 66:337–339

Centella asiatica (Gotu kola) Family: Apiaceae

Medicinal

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in pants of Centella asiatica was reported from Madagascar (Cardin and Moury 2010). The virus-infected gotu kola plants exhibit mosaic, mottling, and yellowish ringspot symptoms in the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

References Cardin L, Moury B (2010) First report of Cucumber mosaic virus in Centella asiatica in Madagascar. J Plant Pathol 92:S4.114

Centranthus ruber (Red valerian) Family: Caprifoliaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

C

478

Centrosema spp.

CMV infection in pants of Centranthus ruber was reported from Southern Italy (Carrieri et al. 2012). The virus-infected red valerian plants show stunting, mild vein mosaic and marked reduction in the size of leaves and flowers, and occasional deformation and reduction of the number of leaves. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sapinoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in pants of Centranthus ruber was reported from Pennsylvania (USA) (Hausbeck et al. 1992). The virus-infected red valerian plants exhibit mottle mosaic, necrosis, ringspots, and chlorosis symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Carrieri R, Sorrentino R, Ragozzino E, Alioto D (2012) Red valerian (Centranthus ruber L.): wild host of Cucumber mosaic virus in uncultivated areas of Campania region (Southern Italy). Phytopathol Mediterr 51:397–401 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among green house ornamentals in Pennsylvania. Plant Dis 76:795–800

Centrosema spp. Family: Fabaceae

Forage crop

African cassava mosaic virus Taxonomic position Genus: Begomovirus

(ACMV)

Family: Geminiviridae

ACMV infection in plants of Centrosema pubescens was reported from the Democratic Republic of Congo (Monde et al. 2010). The virus-infected Centrosema plants exhibit mosaic symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of ACMV, refer to Manihot esculenta.

Bean common mosaic necrosis virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

(BCMNV)

Centrosema spp.

479

BCMNV was reported on Centrosema pubescens in Uganda by Sengooba et al. (1997). The virusinfected Centrosema plants exhibit mild mosaic symptoms. The virus is transmitted by an aphid vector Aphis fabae in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BCMNV, refer to Phaseolus vulgaris.

Centrosema mosaic virus

(CenMV)

Taxonomic position CenMV is a tentative member of the genus Potyvirus and family Potyviridae. Geographical distribution CenMV infection in plants of Centrosema pubescens was reported from Nigeria (Odedara et al. 2007). An earlier report of a virus under the same name from Papua and New Guinea (van Velsen and Crowley, 1961) may be the same or a different virus. Symptoms and host(s) The virus-infected Centrosema plants exhibit symptoms of leaf malformation, mosaic, and mottling. Transmission The virus is probably transmitted by aphid vectors, in a non-persistent manner. The virus is mechanically sap-transmissible. Virion properties and genome The virions are flexuous filaments particles about 750 nm length and 18 nm wide. A partially purified preparation of the Papua and New Guinea isolate had particles of average length 600 nm, or 580 nm if particles of 800 nm were excluded (Crowley and Francki 1963). The genome consists of a single molecule of positive sense single stranded RNA. No sequence information is currently available for either the Nigeran or Papua and New Guinea virus.

Centrosema yellow spot virus Taxonomic position Genus: Begomovirus

(CeYSV)

Family: Geminiviridae

Geographical distribution CeYSV infection in plants of Centrosema brasilianum was reported from Brazil (Silva et al. 2012). Symptoms and host(s) The virus-infected Centrosema plants exhibit mild foliar yellow spot symptoms. Transmission The transmission of CeYSV has not been investigated. It is likely that, in common with other begomoviruses, CeYSV is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner.

C

480

Centrosema spp.

Virion properties and genome The structure of the virions of CeYSV has not been investigated. In common with all geminiviruses, the virions of CeYSV are likely geminate (twinned quasi-icosahedra). Only a single isolate of CeYSV has been characterized. The precise nature of the genome (whether monopartite or bipartite) is unclear since no DNA B component has been reported. The characterized genome/DNA-A genomic component of CeYSV is 2675 nt (JN419002 = NC_016998) (Briddon 2001; Silva et al. 2012; Brown et al. 2015; Zerbini et al. 2017). The genome of CeYSV encodes the five genes typically encoded by monopartite begomoviruses and the DNA-A components of bipartite begomoviruses originating from the New World. The expression and function of the genes have not been investigated.

Cowpea mild mottle virus Taxonomic position Genus: Carlavirus

(CPMMV)

Family: Betaflexiviridae

CPMMV infection in plants of Centrosema spp. was reported from Lome, Togo (West Africa) by Gumedzoe (1993). The virus-infected Centrosema plants exhibit mosaic and mottling symptoms. The virus is transmitted by a whitefly vector, Bemisia tabaci, in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of CPMMV, refer to Vigna unguiculata.

Cowpea severe mosaic virus Taxonomic position Genus: Comovirus

(CPSMV)

Family: Secoviridae

CPSMV infection in plants of Centrosema pubescens was reported from Central Brazil (Lin et al. 1982). The virus-infected Centrosema plants exhibit symptoms of interveinal chlorosis and mosaic on leaves. The virus is transmitted by beetle vectors and also through mechanical sap-inoculation. For more details of CPSMV, refer to Vigna unguiculata.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Centrosema spp. was reported from Guadeloupe and Lome in Togo (West Africa) (Gumedzoe 1993). The virus-infected Centrosema plants exhibit mosaic mottling symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of CMV, refer to Cucumis sativus.

East African cassava mosaic virus Taxonomic position Genus: Begomovirus

(EACMV)

Family: Geminiviridae

Centrosema spp.

481

EACMV infection in plants of Centrosema pubescens was reported from the Democratic Republic of Congo (Monde et al. 2010). The virus-infected Centrosema plants exhibit mosaic symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of EACMV, refer to Manihot esculenta.

Passion fruit woodiness virus Taxonomic position Genus: Potyvirus

(PWV)

Family: Potyviridae

PWV infection in plants of Centrosema pubescens was reported from Australia (Greber 1971). The virus-infected Centrosema plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PWV, refer to Passiflora edulis.

Soybean mosaic virus Taxonomic position Genus: Potyvirus

(SMV)

Family: Potyviridae

SMV infection in Centrosema macrocarpum plants was reported from Columbia (Morales et al. 1990). The virus-infected Centrosema plants exhibit symptoms of mosaic and leaf distortion. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SMV, refer to Glycine max.

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Crowley NC, Francki RIB (1963) Purification and some properties of Centrosema mosaic virus. Aust J Biol Sci 16:468–472 Greber RS (1971) A mosaic disease of Centrosema pubescens Benth. caused by Passionfruit woodiness virus. Queensland J Agric Anim Sci 28:115–119 Gumedzoe MY (1993) Major virus diseases of medicinal plants in West Africa. Acta Hortic (ISHS) 331:307–310 Lin MT, Anjos JRN, Rios GP (1982) Cowpea severe mosaic virus in five legumes in central Brazil. Plant Dis 66:67–70 Monde G, Walangululu J, Winter S, Bragard C (2010) Dual infection by cassava begomoviruses in two leguminous species (Fabaceae) in Yangambi, Northeastern Democratic Republic of Congo. Arch Virol 155:1865–1869 Morales FJ, Niessen AI, Castano M, Calvert L (1990) Detection of a strain of Soybean mosaic virus affecting tropical forage species of Centrosema. Plant Dis 74:648–651 Odedara OO, Hughes JDA, Tarawali SA, Odebode AC, Winter S (2007) Characterisation of a potyvirus from Centrosema pubescens Benth. Trop Sci 47(1):3–15 Sengooba TN, Spence NJ, Walkey DGA, Allen DJ, Femi Lana A (1997) The occurrence of Bean common mosaic necrosis virus in wild and forage legumes in Uganda. Plant Pathol 46:95–103 Silva SJC, Castillo-Urquiza GP, Hora BT Jr, Assuncao IP, Lima GSA, Pio-Ribeiro G, Mizubuti ESG, Zerbini FM (2012) Species diversity, phylogeny and genetic variability of begomovirus populations infecting leguminous weeds in northeastern Brazil. Plant Pathol 61:457–467

C

482

Cerastium spp. (Chickweed)

Van Velsen RJ, Crowley NC (1961) Centrosema mosaic: a plant virus disease transmitted by both aphids and plant bugs. Nature 189:858 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Cerastium spp. (Chickweed) Family: Caryophyllaceae

Weed host

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Cerastium glomeratum was reported from Japan (Okuda et al. 2010). The virus-infected chickweed plants exhibit necrotic spot symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

References Okuda M, Fuji S, Okuda S, Sako K, Iwanami T (2010) Evaluation of the potential of thirty two weed species as infection sources of Impatiens necrotic spot virus. J Plant Pathol 92:357–361

Ceratobium spp. Family: Orchidaceae

Ornamental

Ceratobium mosaic virus Taxonomic position Genus: Potyvirus

(CerMV)

Family: Potyviridae

Geographical distribution CerMV infection in plants of Ceratobium spp. was reported from Australia (Mackenzie et al. 1998). Symptoms and host(s) The virus-infected Ceratobium plants exhibit mosaic symptoms. Transmission The virus is transmitted by aphid vectors and the virus is also mechanically sap-transmissible.

Cercis canadensis (Redbud)

483

Virion properties and genome The virions are non-enveloped, flexuous filaments, and 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA (Revers and Garcia 2015). A partial genome sequence of 1720 nt is available (AF022442) (Mackenzie et al. 1998; Wylie et al. 2017).

C References Mackenzie AM, Nolan M, Wei KJ, Clements MA, Gowanlock D, Wallace BJ, Gibbs AJ (1998) Ceratobium mosaic potyvirus: another virus from orchids. Arch Virol 143:903–914 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Cercis canadensis (Redbud) Family: Fabaceae

Ornamental

Redbud yellow ringspot-associated emaravirus Taxonomic position Genus: Emaravirus

(RYRSaV)

Family: Fimoviridae

Geographical distribution RYRSaV infection in plants of Cercis canadensis was reported from Arkansas, USA (Kim and Martin 1978; Kim et al. 2001; Laney 2010; Laney et al. 2010; Di Bello et al. 2016). Symptoms and host(s) The virus-infected redbud plants exhibit symptoms of vein-clearing, yellow ringspots, and oak leaf pattern on leaves; sometimes tree decline is noticed. Some infected plants are symptomless. No other hosts were reported to be susceptible to the virus. Transmission The virus is transmitted by an eriophyid mite vector (Kim et al. 2001). Eriophyid mite Aculops cercidis was prevalent on the redbud trees but was not found to be a vector of the virus. The virus is transmissible by grafting to redbud and a few faba bean plants which tested positive for the virus and developed symptoms after grafting (Di Bello et al. 2016). Virion properties and genome The virions are quasi-spherical, double membrane-bound particles (DMBs) of 80–100 mm in diameter. The genome is composed of five segments of negative-sense ssRNAs: RNA-1, 7049 nt (JF795479); RNA-2, 2220 nt (JF795480); RNA-3, 1414 nt (JF795481); RNA-4, 1513 nt (JF795482); and RNA-5, 1272 nt (KU904300) (Kim et al. 2001; Di Bello et al. 2016; Elbeaino et al. 2018).

484

Cestrum spp.

References Di Bello PL, Laney AG, Druciarek T, Ho T, Gergerich RC, Keller KE, Martin RR, Tzanetakis IE (2016) A novel Emaravirus is associated with redbud yellow ringspot disease. Virus Res 222:41–47 Elbeaino T, Digiaro M, Mielke-Ehret N, Muelbach H-P, Martelli GP, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Fimoviridae. J Gen Virol 99:1478–1479 Kim K, Martin E (1978) Virus-like particles associated with yellow ringspot of redbud. Phytopathol News 12:119 Kim K, Ahn K, Gergelich R, Kim S-B (2001) Possible etiology of eriophyid mite-borne pathogens associated with double membrane-bound particles. In: Harris KF, Smith OP, Dodds JF (eds) Virus–insect–plant interactions. Academic Press, pp 29–50 Laney AG (2010) Characterization of the causal agents of Rose rosette and Redbud yellow ringspot. MS thesis. University of Arkansas. Laney AG, Gergerich R, Keller K, Martin R, Tzanetakis I (2010) Rose rosette and redbud yellow ringspot are caused by two new emaraviruses. Phytopathology 100:S67

Cestrum spp. Family: Solanaceae

Cestrum ringspot virus

Ornamental

(CeRSV)

Taxonomic position CeRSV is a tentative member of the genus Dichoravirus and family Rhabdoviridae. Geographical distribution CeRSV infection in plants of Cestrum nocturnum was reported from Brazil (Guidotti et al. 2006; Kitajima et al. 2010). Symptoms and host(s) The virus-infected cestrum plants exhibit chlorotic ringspot symptoms. Transmission The virus is transmitted by the mite vectors, Brevipalpus phoenicis and B. obovatus (Guidotti et al. 2006). Virion properties and genome Small rod-like particles of c. 40  100 nm are observed in the nucleus and within the endoplasmic reticulum of infected cells, with little evidence of nuclear viroplasms (Kitajima et al. 2003). The genome is presumed to be composed of two segments of negative-sense single-stranded RNA of c.6.4 kb (RNA 1) and 6 kb (RNA 2), but no genome sequence is available.

Cestrum yellow leaf curling virus

(CmYLCV)

Synonyms Cestrum virus Taxonomic position Genus: Soymovirus

Family: Caulimoviridae

Cestrum spp.

485

Geographical distribution CmYLCV infection in plants of Cestrum parqui was reported from Italy (Stavolone et al. 2003). Symptoms and host(s) The virus-infected cestrum plants exhibit mosaic, leaf curling, and stunting symptoms.

C Transmission The virus is mechanically sap-transmissible. The experimental host range consists of Cestrum parqui, Cestrum elegans, and Nicotiana clevelandii (all in the family Solanaceae). Virion properties and genome The virions are isometric, non-enveloped, and 50 nm in diameter with no obvious surface structure. The genome is a monopartite, circular double-stranded DNA of 8253 bp (NC_004324) (Stavolone et al. 2003; Hibi 2011).

Citrus exocortis viroid Taxonomic position Genus: Pospiviroid

(CEVd)

Family: Pospiviroidae

CEVd was detected from Cestrum auriculatum in Italy (Luigi et al. 2011a, b). The viroid was detected from a symptomless infection in C. auriculatum. The viroid is mechanically sap-transmissible. For more details of CEVd, refer to Citrus spp.

Clerodendrum chlorotic spot dichorhavirus Taxonomic position Genus: Dichorhavirus

(ClCSV)

Family: Rhabdoviridae

A plant of Cestrum nocturnum growing near a ClCSV-infected Clerodendrum  speciosum in Brazil was found to have chlorotic spots on the leaves; the causal virus was determined to be ClCSV (Kitajima et al. 2008). The virus is transmitted by mite vectors and also by mechanical sap-inoculation. For more details of ClCSV, refer to Clerodendrum spp.

Pedilanthus leaf curl virus Taxonomic position Genus: Begomovirus

(PeLCV)

Family: Geminiviridae

PeLCV infection in plants of night blooming jasmine (Cestrum nocturnum) was reported from India (Srivastava et al. 2014). The virus-infected blooming jasmine plants exhibit leaf curl symptoms. The

486

Cestrum spp.

virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of PeLCV, refer to Pedilanthus tithymaloides.

Potato spindle tuber viroid Taxonomic position Genus: Pospiviroid

(PSTVd)

Family: Pospiviroidae

PSTVd infection in plants of Cestrum spp. (C. auricantum, C. rubrum, C. x cultum, C. endlicheri, and C. nocturnum) was reported from Italy (Luigi et al. 2011a, b). The viroid-infected cestrum plants display no symptoms. The viroid is mechanically sap-transmissible. For more details of PSTVd, refer to Solanum tuberosum.

Tomato apical stunt viroid Taxonomic position Genus: Pospiviroid

(TASVd)

Family: Pospiviroidae

TASVd infection in plants of Cestrum spp. was reported from the Netherlands (Verhoeven et al. 2008, 2012). The viroid-infected cestrum plants are symptomless. The viroid is mechanically sap-transmissible. For more details of TASVd, refer to Solanum lycopersicum.

References Guidotti AR, Rodrigues V, Machado MA, Freitas-Astua J (2006) Transmission of the Cestrum ringspot virus by different active phases of the mite Brevipalpus obovatus. Summa Phytopathol 32(S):75 Hibi T (2011) Soymovirus. Caulimoviridae. In: The Springer index of Viruses. Springer, New York, pp 287–292. https:// doi.org/10.1007/978-0-387-95919-1_44 Kitajima EW, Chagas CM, Rodrigues JCV (2003) Brevipalpus-transmitted plant vírus and virus-like diseases: cytopathology and some recent cases. Exp Appl Acarol 30:135–160 Kitajima EW, Kubo KS, Ferreira PTO, Alc^antara BK, Boari AJ, Gomes RT, Freitas-Astúa J, Rezende JAM, Morais GJ, Salaroli RB (2008) Chlorotic spots on Clerodendrum, a disease caused by a nuclear type of Brevipalpus (Acari: Tenuipalpidae) transmitted virus. Sci Agric 65:36–49 Kitajima EW, Rodrigues JCV, Freitas-Astua J (2010) An annotated list of ornamentals naturally found infected by Brevipalpus mite-transmitted viruses. Sci Agric (Piracicaba, Braz) 67:348–371 Luigi M, Luison D, Tomassoli L, Faggioli F (2011a) Natural spread and molecular analysis of pospiviroids infecting ornamentals in Italy. J Plant Pathol 93:491–495 Luigi M, Luison D, Tomassoli L, Faggioli F (2011b) First report of Potato spindle tuber and Citrus exocortis viroids in Cestrum spp. in Italy. New Dis Rep 23:4 Srivastava A, Kumar S, Raj SK (2014) Association of Pedilanthus leaf curl virus with yellow mottling and leaf curl symptoms in two jasmine species grown in India. J Gen Plant Pathol 80:370–373 Stavolone L, Ragozzino A, Hohn T (2003) Characterization of Cestrum yellow leaf curling virus: a new member of the family Caulimoviridae. J Gen Virol 84:3459–3464 Verhoeven JTJ, Jansen CCC, Roenhorst JW (2008) First report of pospiviroids infecting ornamentals in the Netherlands: Citrus exocortis viroid in Verbena sp., Potato spindle tuber viroid in Brugmansia suaveolens and Solanum jasminoides, and Tomato apical stunt viroid in Cestrum sp. Plant Pathol 57:399 Verhoeven JTJ, Botermans M, Meekes ETM, Roenhorst JW (2012) Tomato apical stunt viroid in the Netherlands: most prevalent Pospiviroid in ornamentals and first outbreak in tomatoes. Eur J Plant Pathol 133:803–810

Cheesemania radicata

487

Chamaescilla corymbosa (Blue stars, Blue squill) Family: Asparagaceae

Ornamental

Blue squill virus A

(BSVA)

Taxonomic position Genus: Potyvirus

C

Family: Potyviridae

Geographical distribution BSVA infection in plants of Chamaescilla corymbosa was reported from Australia (Wylie et al. 2012). Symptoms and host(s) The virus-infected blue squill plants exhibit faint chlorotic mottle symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner, and the virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments particles, and 750 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9842 nt (JQ807999 = NC_019415) (Wylie et al. 2012; Revers and Garcia 2015; Wylie et al. 2017).

References Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Wylie SJ, Luo H, Li H, Jones MG (2012) Multiple polyadenylated RNA viruses detected in pooled cultivated and wild plant samples. Arch Virol 157(2):271–284 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Cheesemania radicata Family: Brassicaceae

Ornamental

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

BWYV infection in plants of Cheesemania spp. was reported from Australia (Guy 1987). The virusinfected Cheesemania plants exhibit symptoms of chlorotic blotching in the older leaves. The virus is

488

Cheiranthus spp. (Wallflower)

transmitted by aphid vectors in a circulative and non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of BWYV, refer to Beta vulgaris.

References Guy PL (1987) Beet western yellows virus infection of Cardamine spp. and Cheesemania radicata. Australas Plant Pathol 16:43–44

Cheiranthus spp. (Wallflower) Synonyms Erysimum spp. Family: Brassicaceae

Erysimum latent virus Taxonomic position Genus: Tymovirus

Ornamental

(ErLV)

Family: Tymoviridae

Geographical distribution ErLV-infected Erysimum spp. was reported from Germany (Shukla and Schmelzer 1972). Symptoms and host(s) The virus-infected wallflower plants do not exhibit any symptoms. Transmission The virus is transmitted by the flea beetle vector Phyllotreta spp. in a semi-persistent manner. The virus is transmissible by mechanical sap-inoculation to more than nine plant families. The virus is not transmissible by contact between plants, not transmissible by seed, and not transmissible by pollen. Virion properties and genome The virions are non-enveloped, isometric capsid with T = 3 icosahedral symmetry, about 30 nm in diameter. The genome consists of a single molecule of positive-sense, ssRNA of 6035 nt (AF098523 = NC_001977). The 30 -terminus has a tRNA-like structure (Srifah et al. 1992; Martelli et al. 2002).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV was reported from Cheiranthus cheiri in Iran, causing necrotic leaf spot, chlorosis, and stunting and reducing both yield and quality (Ghotbi 2013). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Chelidonium majus (Greater celandine, Tetterwort)

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

489

(TSWV)

Family: Tospoviridae

TSWV infection occurs wherever Cheiranthus cheiri plants are grown (Smith 1957). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sapinoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Cheiranthus cheiri was reported from the USA, New Zealand, Europe, and Iran (Chamberlain 1954; Bahar et al. 1984; Farzadfar et al. 2005; Bellardi et al. 2013). The virusinfected wallflower plants exhibit dark mottle, stripes, leaf yellows, and malformation symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TuMV, refer to Brassica rapa.

References Bahar M, Danesh D, Dehghan M (1984) Turnip mosaic virus in wallflower. Iran J Plant Pathol 21:33–39 Bellardi MG, Cavicchi L, De Stradis A, Panno S, Davino S (2013) Molecular characterization and phylogenetic analysis of Turnip mosaic virus (TuMV) in Erysimum linifolium L. in Italy. Int Res J Pl Sci 4:97–102 Chamberlain EE (1954) Plant viruses in New Zealand. N.Z. DSIR Bull 108:153–160 Farzadfar SH, Ohshima K, Pourrahim R, Golnaraghi AR, Jalali S, Ahoonmanesh A (2005) Occurrence of Turnip mosaic virus on ornamental crops in Iran. Plant Pathol 54:261 Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iran J Plant Pathol 49:41 Martelli GP, Sabanadzovic S, Abou-Ghanem Sabanadzovic N, Edwards MC, Dreher T (2002) The family Tymoviridae. Arch Virol 147(9):1837–1846 Shukla DD, Schmelzer K (1972) Studies on viruses and virus diseases of cruciferous plants. IV. The previously undescribed Erysimum latent virus. Acta Phytopath Acad Sci Hung 7:157–167 Smith KM (1957) A textbook of plant virus diseases, 2nd edn. Churchill, London, 572 pp Srifah P, Keese P, Weiller G, Gibbs A (1992) Comparisons of the genomic sequences of Erysimum latent virus and other tymoviruses: a search for the molecular basis of their host specificities. J Gen Virol 73:1437–1447

Chelidonium majus (Greater celandine, Tetterwort) Family: Papaveraceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Medicinal plant

(CMV)

Family: Bromoviridae

C

490

Chelidonium majus (Greater celandine, Tetterwort)

CMV infection in plants of Chelidonium majus was reported from Czechoslovakia, Switzerland, and Moldova (Dashkeeva et al. 1966; Hani 1971; Brcak 1979). The virus-infected tetterwort plants exhibit mild mottling symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Tetterwort vein chlorosis virus Taxonomic position Genus: Crinivirus

(TVCV)

Family: Closteroviridae

Geographical distribution TVCV reported infecting Chelidonium majus from Korea (Zhao et al. 2015).

Symptoms and host(s) The virus-infected tetterwort plants show veinal chlorosis and leaf distortion symptoms.

Transmission The virus is transmitted by whitefly vectors in a semi-persistent manner.

Virion properties and genome The virions are non-enveloped, bipartite filamentous particles about 650–850 nm and 700–900 nm in length, and 10–13 nm in diameter. The genome is bipartite single-stranded RNA; that of isolate Yesan that consists of an 8467 nt RNA1 (KR002686) and an 8113 nt RNA2 (KR002687). RNA1 consists of four open reading frames (ORFs) designated as ORF1a (nt 67–6045), ORF1b (nt 6044–7561), P6a (ORF2, nt 7577–7738), and P19 (ORF3, nt 7741–8232). RNA2 consists of nine ORFs coding for p9a (nt 125–358), p5 (nt 526–660), the heat shock protein 70 homolog (HSP70h, nt 1331–3004), p6b (nt 3005–3169), p60 (nt 3163–4716), p9b (nt 4698–4944), p23 (major coat protein, nt 5055–5780), p54 (minor coat protein, nt 5780–7204), and p27 (nt 7211–7900) (Kreuze 2011; Zhao et al. 2015).

References Brcak J (1979) Isolates of Cucumber mosaic virus from spontaneously infected plants of Chelidonium majus and Impatiens parviflora. Biol Plant 21:220–223 Dashkeeva KN, Spatarenko SS, Vetrova FM (1966) On the role of weeds in persistence and spread of the Cucumber mosaic virus. Infektsionnye Zabolevaniya kul’turnykh Rasteniï Moldavii (Kishinev) 6:22–27 Hani A (1971) Zur Epidemiologie des Gurkenmosaikvirus im Tessin. Phytopathol Z 72:115–144 Kreuze JF (2011) Crinivirus. Closteroviridae. In: The Springer index of Viruses. Springer, New York, pp 335–342. https:// doi.org/10.1007/978-0-387-95919-1_51 Zhao F, Yoo RH, Lim S, Igori D, Lee S-H, Moon JS (2015) Nucleotide sequence and genome organization of a new proposed crinivirus, Tetterwort vein chlorosis virus. Arch Virol 160:2899

Chenopodium spp.

491

Chenopodium spp. Family: Chenopodiaceae

Leafy vegetable

Alfalfa mosaic virus

(AMV)

Taxonomic position Genus: Alfamovirus

Family: Bromoviridae

AMV infection in plants of Chenopodium spp. was reported from Saudi Arabia (Al-Shahwan et al. 2017). The virus-infected Chenopodium plants exhibit yellow mosaic on the leaves. The virus is transmitted by a number of aphid species in a non-persistent manner, and also through mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Chenopodium spp. was reported from Italy (Vicchi and Bellardi 1988). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Beet curly top virus Taxonomic position Genus: Curtovirus

(BCTV)

Family: Geminiviridae

BCTV infection in plants of Chenopodium album and C. murale was reported from California (USA) (Creamer et al. 1996). The virus is transmitted by a leafhopper vector Circulifer tenellus in a persistent (circulative, non-propagative) manner. The virus is not mechanically sap-transmissible. For more details of BCTV, refer to Beta vulgaris.

Beet chlorosis virus Taxonomic position Genus: Polerovirus

(BChV)

Family: Luteoviridae

BChV infection in plants of Chenopodium capitatum was reported from France (Hauser et al. 2000). The virus is transmitted by aphid vectors in a circulative and non-propagative manner. The virus is not mechanically sap-transmissible. For more details of BChV, refer to Beta vulgaris.

C

492

Chenopodium spp.

Chenopodium leaf curl virus Taxonomic position Genus: Begomovirus

(ChLCV)

Family: Geminiviridae

Geographical distribution ChLCV infection in plants of Chenopodium ambrosioides was reported from Florida (USA) (Ng et al. 2011). Symptoms and host(s) The virus-infected Chenopodium plants exhibit leaf curling symptoms. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2626 nt (HM626515) (Briddon 2001; Ng et al. 2011; Brown et al. 2015; Zerbini et al. 2017).

Chenopodium necrosis virus Taxonomic position Genus: Unassigned

(ChNV)

Family: Tombusviridae

Geographical distribution ChNV infection in plants of Chenopodium spp. was reported from England (Tomlinson et al. 1983). Symptoms and host(s) The virus-infected Chenopodium plants exhibit systemic necrosis symptoms. Transmission The virus is transmitted by a fungal vector Olpidium brassicae. The virus is mechanically saptransmissible. In Chenopodium spp. the virus induced local lesions and systemic infection. Virion properties and genome The virions are isometric and icosahedral (T = 3), non-enveloped, and 28–34 nm in diameter. The genome is a monopartite, linear single-stranded positive-sense RNA.

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(CABYV)

Family: Luteoviridae

CABYV infection in plants of Chenopodium murale was reported from Saudi Arabia (Al-Saleh et al. 2015). The virus-infected plants exhibit interveinal chlorosis and yellowing symptoms. The virus is

Chenopodium spp.

493

transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of CABYV, refer to Cucumis melo.

Grapevine Pinot gris virus

(GPGV)

C

Taxonomic position Genus: Trichovirus

Family: Betaflexiviridae

GPGV infection in plants of Chenopodium album was reported from Italy (Gualandri et al. 2017). The virus is mechanically sap-transmissible. For more details of GPGV, refer to Vitis vinifera.

Peanut stunt virus

(PSV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

PSV infection in plants of Chenopodium album was reported from Iran (Amid-Motlagh et al. 2017). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation to a large number of hosts. For more details of PSV, refer to Arachis hypogaea.

Pepino mosaic virus Taxonomic position Genus: Potexvirus

(PepMV)

Family: Alphaflexiviridae

PepMV infection in plants of Chenopodium murale was reported from Spain (Cordoba et al. 2004). The virus is mechanically sap-transmissible. For more details of PepMV, refer to Solanum muricatum.

Potato virus S Taxonomic position Genus: Carlavirus

(PVS)

Family: Betaflexiviridae

PVS infection in plants of Chenopodium botrytis and C. album was reported from Iran (Hosseini and Salari 2017). The virus is transmitted by aphid vectors in a non-persistent manner, and also through mechanical sap-inoculation. For more details of PVS, refer to Solanum tuberosum.

Sowbane mosaic virus Synonyms Rubus chlorotic mottle virus

(SoMV)

494

Taxonomic position Genus: Sobemovirus

Chenopodium spp.

Family: Solemoviridae

Geographical distribution SoMV was first reported in plants of Chenopodium murale from Riverside, California, USA. The virus also occurs in Canada, Central and South America, Europe, Japan, New Zealand, Australia, Bulgaria, Hungary, the former Czechoslovakia, Italy, Japan, and the former Yugoslavia (Teakle 1968; Horvath et al. 1993; Pearson et al. 2006; Eastwell et al. 2010). Symptoms and host(s) The virus-infected Chenopodium plants exhibit systemic chlorotic mottling symptoms. Transmission The virus has been reported to be transmitted by insects belonging to different orders (Diptera, Hemiptera, Thysanoptera), but this is probably a non-specific mechanical transmission by the mouthparts of different insects. The most common vectors are different species of beetle that transmit the virus in a semi-persistent manner. The virus is pollen-borne, and the injuries caused by thrips feeding (Thrips tabaci) will help in the entry of the virus carried in/on the infected pollen (Hardy and Teakle 1992). The virus is transmissible by grafting and also through seed of Chenopodium spp. up to 70% (Bennett and Costa 1961; Dias and Waterworth 1967). Virion properties and genome The virions are isometric, non-enveloped, and 30 nm in diameter. The genome is a polycistronic positive-sense single-stranded RNA (ssRNA) of 3983 nt (AM940437 = NC_011187; GQ845002). The genome contains two overlapping ORFs, ORF2a, and ORF2b (Kado and Black 1968; Hull 1988; Somera et al. 2015).

Squash mosaic virus Taxonomic position Genus: Comovirus

(SqMV)

Family: Secoviridae

SqMV infection in plants of Chenopodium spp. was reported from Morocco and Africa (Lockhart et al. 1982, 1985). The virus-infected Chenopodium plants exhibit mosaic symptoms. The virus is transmitted by a beetle vector Epilachna chrysomelina and also by mechanical sap-inoculation. The virus is seed-transmitted in Chenopodium spp. up to 23% (Lockhart et al. 1985). For more details of SqMV, refer to Cucurbita pepo.

Tomato mosaic virus Taxonomic position Genus: Tobamovirus

(ToMV)

Family: Virgaviridae

ToMV infection in plants of Chenopodium murale was reported from Iran (Ahoonmanesh et al. 1992). The virus-infected Chenopodium plants exhibit symptoms of mosaic and reduced growth. No vector is

Chenopodium spp.

495

involved in the spread of this virus. This virus is transmissible by mechanical sap-inoculation, grafting, and contact between plants. For more details of ToMV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

C Family: Tospoviridae

TSWV infection in plants of Chenopodium album plants was reported from Spain and Italy (Moreno et al. 2004; Grieco et al. 2000). The virus-infected Chenopodium plants exhibit severe mosaic symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Chenopodium murale was reported from Spain and Cyprus (Jorda et al. 2001; Papayiannis et al. 2011). The virus-infected Chenopodium plants exhibit symptomless infections. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The virus is transmissible by grafting but not by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

References Ahoonmanesh A, Bahar M, Ghobadi C (1992) Tomato mosaic virus in Iran. Iran J Plant Pathol 28:1–4 Al-Saleh MA, Al-Shahwan IM, Amer MA, Shakeel MT, Kamran A, Xanthis CK, Orfanidou CG, Katis NI (2015) First report of Cucurbit aphid-borne yellows virus in cucurbit crops in Saudi Arabia. Plant Dis 99:894 Al-Shahwan IM, Abdalla OA, Al-Saleh MA, Amer MA (2017) Detection of new viruses in alfalfa, weeds and cultivated plants growing adjacent to alfalfa fields in Saudi Arabia. Saudi J Biol Sci 24(6):1336–1343 Amid-Motlagh MH, Massumi H, Heydarnejad J, Mehrvar M, Reza Hajimorad M (2017) Nucleotide sequence analyses of coat protein gene of Peanut stunt virus isolates from alfalfa and different hosts show a new tentative subgroup from Iran. Virus Dis 28:295–302 Bennett CW, Costa AS (1961) Sowbane mosaic caused by a seed transmitted virus. Phytopathology 51:546–550 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Cordoba MC, Martinez-Priego LI, Jorda C (2004) New natural hosts of Pepino mosaic virus in Spain. Plant Dis 88:906 Creamer R, Lugue-Williams M, Howo M (1996) Epidemiology and incidence of Beet curly top geminivirus in naturally infected weed hosts. Plant Dis 80:533–535 Dias HF, Waterworth HE (1967) The identification of a seed-borne mosaic virus of Chenopodium amaranticolor and C. quinoa. Can J Bot 45:1285–1295

496

Chionanthus virginicus (White fringe tree)

Eastwell KC, Villamor DV, McKinney CL (2010) Characterization of an isolate of Sowbane mosaic virus. Arch Virol 155:2065–2067 Grieco PD, Conte D, Munno I, Nuzzaci M, de Stradis A (2000) Tomato spotted wilt virus (TSWV) on weeds and wild plants in Metapontino, Basilicata. Informatore Fitopatologico 50:43–46 Gualandri V, Asquini E, Bianchedi P, Covelli L, Brilli M, Malossini U, Bragagna P, Saldarelli P, Si-Ammour A (2017) Identification of herbaceous hosts of the Grapevine Pinot gris virus (GPGV). Eur J Plant Pathol 147:21–25 Hardy VG, Teakle DS (1992) Transmission of sowbane mosaic virus by Thrips tabaci in the presence and absence of virus-carrying pollen. Ann Appl Biol 121:315–320 Hauser S, Stevens M, Mougel C, Smith HG, Fritsch C, Herrback E, Lemaire O (2000) Biological, serological and molecular variability suggest three distinct polerovirus species infecting beet or rape. Phytopathology 90:460–466 Horvath J, Juretic N, Wolf I, Pinter CS (1993) Natural occurrence of Sowbane mosaic virus on Chenopodium hybridum L. in Hungary. Acta Phytopathol Entomol Hung 28:379–389 Hosseini SA, Salari K (2017) Detection and molecular characterisation of Potato virus S of weed reservoirs in Iran. Arch Phytopathol Plant Protect 50:828–838 Hull T (1988) The sobemovirus group. In: Koenig R (ed) The plant viruses. Polyhedral virions with monopartite RNA genomes, vol 3. Plenum Press, New York, pp 113–146 Jorda C, Font I, Martinez P, Juarez M, Ortega A, Lacasa A (2001) Current status and new natural hosts of Tomato yellow leaf curl virus (TYLCV) in Spain. Plant Dis 85:445 Kado CI, Black DR (1968) The molecular weight of Sowbane mosaic virus. Virology 36:137–139 Lockhart BEL, Ferji Z, Hafidi B (1982) Squash mosaic virus in Morrocco. Plant Dis 66:1191–1193 Lockhart BEL, Jebbour F, Lennon AM (1985) Seed transmission of Squash mosaic virus in Chenopodium spp. Plant Dis 69:946–947 Moreno A, de Blas C, Biurrun R, Nebreda M, Palacios I, Duque M, Fereres A (2004) The incidence and distribution of viruses infecting cultivated lettuce, cultivated brassica and associated natural vegetation in Spain. Ann Appl Biol 144:339–346 Ng TF, Duffy S, Polston JE, Bixby E, Vallad GE, Breitbart M (2011) Exploring the diversity of plant DNA viruses and their satellites using vector-enabled metagenomics on whiteflies. PLoS One 6(4):E19050 Papayiannis L, Katis NI, Idris AM, Brown JK (2011) Identification of weed hosts of Tomato yellow leaf curl virus in Cyprus. Plant Dis 95:120–125 Pearson MN, Clover GRG, Guy PL, Fletcher JD, Beever RE (2006) A review of the plant virus, viroid and mollicute records for New Zealand. Aust Plant Pathol 35:217–252 Somera M, Sarmiento C, Truve E (2015) Overview on sobemoviruses and a proposal for the creation of the family Sobemoviridae. Viruses 7:3076–3115 Teakle DS (1968) Sowbane mosaic virus infecting Chenopodium trigonon in Queensland. Aust J Biol Sci 21:649–653 Tomlinson JA, Faithfull EM, Webb MJW, Fraser RSS, Seeley ND (1983) Chenopodium necrosis: a distinctive strain of Tobacco necrosis virus isolated from river water. Ann Appl Biol 102:135–147 Vicchi V, Bellardi MG (1988) The role of weeds in the epidemiology of gladiolus viruses and MLO. Acta Hortic 234:371–378 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Chionanthus virginicus (White fringe tree) Family: Oleaceae

Tobacco rattle virus Taxonomic position Genus: Tobravirus

Medicinal

(TRV)

Family: Virgaviridae

TRV infection in plants of Chionanthus virginica was reported from East Germany (Cooper 1979; Schmelzer 1970). The virus-infected white fringe tree plants exhibit yellow ring symptoms. The

Chloris gayana (Rhodes grass)

497

virus is transmissible by mechanical sap-inoculation, and in nature is transmitted by nematode species in the genera Paratrichodorus and Trichodorus. For more details of TRV, refer to Nicotiana tabacum.

References

C

Cooper JI (1979) Virus disease of trees and shrubs. Institute of Terrestrial Ecology, Cambridge, 74 pp Schmelzer K (1970) Studies on viruses of ornamental and wild woody plants. 7th part: additional reports on Buddleia, Viburnum, Caryopteris and Philadelphus and viruses on Leycesteria, Chionanthus, Ribes, Hydrangea, Syringa, Spiraea and Catalpa. Phytopathol Z 67:285–326

Chirita spp. Family: Gesneriaceae

Ornamental

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV infection in plants of Chirita spp. was reported from the USA (Zettler and Nagel 1983). The symptoms on the foliage of infected plants were inconspicuous. There is no known vector for this virus. The virus is mechanically sap-transmissible and also transmissible by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

References Zettler FW, Nagel J (1983) Infection of cultivated gesneriads by two strains of Tobacco mosaic virus. Plant Dis 67:1123–1125

Chloris gayana (Rhodes grass) Family: Poaceae

Forage crop

Chloris striate mosaic virus Taxonomic position Genu: Mastrevirus

(CSMV)

Family: Geminiviridae

Geographical distribution CSMV infection was first reported in plants of Chloris gayana from Australia by Grylls (1963). The virus has only been identified in Australia (Francki et al. 1979; Francki and Hatta 1980; Andersen et al. 1988).

498

Chloris gayana (Rhodes grass)

Symptoms and host(s) The virus-infected plants of Rhodes grass exhibit chlorotic streaking along the leaves. The natural hosts of CSMV are C. gayana, Brachiaria subquadipara, Dactyloctenium australe, Dactylis glomerata, Ixophorus unisetus, Microlaena stipoides, Paspalum dilatatum, Avena sativa (oats), Hordeum vulgare (barley), Triticum aestivum (wheat), and Zea mays (maize) (Greber 1989). Transmission The virus is transmitted by the leafhopper vectors Nesoclutha pallida and Nesoclutha obscura (Greber 1989). Transmission was achieved with a minimum transmission access period of 12 h on wheat. The latent period between acquisition and transmission was found to be about 6 h. This is consistent with a circulative, non-propagative manner of transmission. The virus is not transmissible by mechanical inoculation. The virus is not transmissible by seed. Virion properties and genome The virions of CSMV are geminated (twinned quasi-isometric), 18  30 nm (Hatta and Francki 1979; Francki et al. 1979). Based on electron micrographs of CSMV particles, Hatta and Francki (1979) produced a model of the structure of the geminate particle in which the structure consists of two T-1 incomplete icosahedra formed by 22 “morphological units” (capsomeres). The two halves are joined where there a missing capsomer, forming a distinct waist to the particle. This model corresponds well to the more recently produced structures for Maize streak virus and African cassava mosaic virus. Francki et al. (1980) showed the genome of CSMV to consist of a circular, single-stranded DNA, with each geminate particle containing one molecule. The genome of CSMV comprises a single circular single-stranded DNA of ~2750 nt (M20021 = NC_001466) (Francki et al. 1979; Andersen et al. 1988; Palmer and Rybicki 1998; Boulton and Davies 2011; Muhire et al. 2013; Zerbini et al. 2017). In common with all mastreviruses, CpCDV is predicted to encode four gene products, two in each orientation. Based on sequence homology to previously studied mastreviruses, these would code for the coat protein and movement protein in the virion-sense and in the complementary-sense for the replication-associated protein (Rep) and Rep A.

References Andersen MT, Richardson KA, Harbison S-A, Morris BAM (1988) Nucleotide sequence of the geminivirus Chloris striate mosaic virus. Virology 164:443–449 Boulton MI, Davies JW (2011) Mastrevirus. Geminiviridae. In: The Springer index of Viruses. Springer, New York, pp 597–603. https://doi.org/10.1007/978-0-387-95919-1_83 Francki RIB, Hatta T (1980) Chloris striate mosaic virus. Descriptions of plant viruses, No. 221. Commonwealth Mycological Institute and Association of Applied Biologists, Kew Francki RIB, Hatta T, Grylls NE, Grivell CJ (1979) The particle morphology and some other properties of Chloris striate mosaic virus. Ann Appl Biol 91:51–59 Francki RIB, Hatta T, Boccardo G, Randles JW (1980) The composition of Chloris striate mosaic virus, a geminivirus. Virology 101:233–241 Greber RS (1989) Biological characteristics of grass geminiviruses from eastern Australia. Ann Appl Biol 114:471–480 Grylls NE (1963) A striate mosaic virus disease of grasses and cereals in Australia transmitted by the cicadellid Nesoclutha obscura. Aust J Agric Res 14:143–152 Hatta T, Francki RIB (1979) The fine structure of Chloris striate mosaic virus. Virology 92:428–435 Muhire B, Martin DP, Brown JK, Navas-Castillo J, Moriones E, Zerbini FM, Rivera-Bustamante R, Malathi VG, Briddon RW, Varsani A (2013) A genome-wide pairwise-identity-based proposal for the classification of viruses in the genus Mastrevirus (family Geminiviridae). Arch Virol 158(6):1411–1424 Palmer KE, Rybicki EP (1998) The molecular biology of mastreviruses. Adv Virus Res 50:183–234 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Chrozophora tinctoria (Dyer’s croton)

499

Choisya ternata (Mexican orange blossom) Family: Rutaceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

C

Family: Secoviridae

ArMV was reported from the UK in container-grown plants of Choisya ternata showing stunting, together with apical dieback of young shoots; the virus was most readily detected in the roots (Mumford et al. 2002). This virus is transmitted by nematode vectors (Xiphinema spp.) in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

References Mumford RA, Blockley AL, Jarvis B, Wright DM (2002) Isolation of Arabis mosaic virus from Mexican orange blossom (Choisya ternata) in the UK. Plant Pathol 51:395

Chrozophora tinctoria (Dyer’s croton) Family: Euphorbiaceae

Weed host

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

CGMMV infection in plants of Chrozophora tinctoria was reported from Israel (Shargil et al. 2017). The virus is mechanically sap-transmissible and also transmitted through contact between plants. The virus is transmissible through soil and irrigation water contaminated with infected plant debris. The virus is also pollen transmissible. For more details of CGMMV, refer to Cucumis sativus.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Chrozophora tinctoria was reported from Cyprus (Papayiannis et al. 2011). The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. The virus is transmitted by grafting but not transmitted by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

500

Chrysanthemum spp.

References Papayiannis L, Katis NI, Idris AM, Brown JK (2011) Identification of weed hosts of Tomato yellow leaf curl virus in Cyprus. Plant Dis 95:120–125 Shargil D, Smith E, Lachman O, Reingold V, Darzi E, Tam Y, Dombrovsky A (2017) New weed hosts for Cucumber green mottle mosaic virus in wild Mediterranean vegetation. Eur J Plant Pathol 148:473–480

Chrysanthemum spp. Synonyms

Dendranthema spp. Family: Asteraceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

Family: Secoviridae

ArMV infection in plants of Chrysanthemum spp. was reported from Iran (Ghotbi and Shahraeen 2005). The virus-infected chrysanthemum plants exhibit symptoms of mosaic, leaf chlorosis, small necrotic lesions, and leaf malformation and deformation. This virus is transmitted by nematode vectors (Xiphinema spp.) in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Bidens mottle virus Taxonomic position Genus: Potyvirus

(BiMoV)

Family: Potyviridae

BiMoV infection in plants of Chrysanthemum coronarium was reported from Taiwan (Chen and Lee 2012). The virus-infected chrysanthemum plants show foliar mosaic, deformation, and stunting symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BiMoV, refer to Bidens spp.

Carnation Italian ringspot virus Taxonomic position Genus: Tombusvirus

(CIRV)

Family: Tombusviridae

CIRV infection in plants of Chrysanthemum zawadskii var. latilobum was reported from Korea (Seo et al. 2015). The virus-infected chrysanthemum plants exhibit mosaic and chlorosis symptoms. The

Chrysanthemum spp.

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virus is mechanically sap-transmissible. The virus is graft-transmissible, and use of infected budwood helps spread this virus. There is no known vector for this virus. For more details of CIRV, refer to Dianthus caryophyllus.

Chilli veinal mottle virus

(ChiVMV)

C

Synonyms Chrysanthemum potyvirus Taxonomic position Genus: Potyvirus

Family: Potyviridae

ChiVMV infection in plants of Chrysanthemum spp. was reported from India (Mehra et al. 2009). The virus-infected chrysanthemum plants exhibit chlorotic spotting symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation, and grafting. For more details of ChiVMV, refer to Capsicum annuum.

Chrysanthemum chlorotic mottle viroid Taxonomic position Genus: Pelamoviroid

(CChMVd)

Family: Avsunviroidae

Geographical distribution CChMVd disease occurs wherever Chrysanthemum spp. are grown (Horst et al. 1977; Singh et al. 1978; Yamamoto and Sano 2005; Chung et al. 2006; Zhang et al. 2011a; Cho et al. 2013; AdkarPurushothama et al. 2015; Zhao et al. 2015; Adkar-Purushothama et al. 2017). Symptoms and host(s) The virus-infected chrysanthemum plants exhibit symptoms of yellow-green mottling eventually developing into pronounced general chlorosis with some dwarfing in some cultivars including ‘Yellow Delaware.’ ‘Knob Hill’ exhibited chlorotic spots, vein-clearing, and mild chlorosis, whereas some cultivars, although infected, remained symptomless. Additional symptoms include reduction of leaves, flowers, and the entire plant and a delay in flower development (Dimock et al. 1971). Transmission The virus is mechanically sap-transmissible and host range is confined to chrysanthemum. The viroid is transmissible by grafting and tissue implantation. No insect vector is known (Horst 1987). The virus could be disseminated over long distances in cuttings and other vegetative plants for planting. Etiology and genome properties The genome consists of a single-stranded circular RNA of 399 nt (Y14700 = NC_003540). They are single-stranded covalent circles and have intramolecular base pairing and lack a central conserved region (Navarro and Flores 1997; Flores et al. 2000; De la Pena and Flores 2002; Gora-Sochacka 2004; Giguere et al. 2014a; Cho et al. 2013; Di Serio et al. 2018).

502

Chrysanthemum spp.

Chrysanthemum stem necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(CSNV)

Family: Tospoviridae

Geographical distribution CSNV infection in plants of Chrysanthemum spp. was reported from South America, Brazil, Netherlands, Slovenia, the UK, New Zealand, Japan, Korea, and Belgium (Nagata et al. 1994; Bezerra et al. 1996, 1999; Verhoeven et al. 1996; Ravnikar et al. 2003; Mumford et al. 2003; CABI/EPPO 2005; EPPO 2006; Matsuura et al. 2007; Takeshita et al. 2011; De Jonghe et al. 2013; Yoon et al. 2017). Symptoms and host(s) The virus-infected chrysanthemum plants exhibit symptoms of mild or severe necrotic streaks on the stem, wilting of leaves and stems, and chlorotic or necrotic spots and rings on some leaves. However, symptoms of CSNV may be more severe and can result in complete necrosis of the stem resulting in wilting of sections of plants. Symptoms in some chrysanthemum cultivars appear as necrotic lesions surrounded by yellow areas on leaves followed by necrosis on stems, peduncles, and floral receptacles. Transmission The virus is transmitted and spread in nature by thrips vectors Frankliniella occidentalis and F. schultzei in a persistent-propagative manner (Bezerra et al. 1999; Nagata and de Avila 2000; Nagata et al. 2004; Riley et al. 2011; Okuda et al. 2013). The virus is mechanically sap-transmissible and infects Chrysanthemum spp., Lisianthus spp., and tomato. The virus could be disseminated over long distances in cuttings and other vegetative plants for planting. CSNV is unlikely to be seed-transmitted. Virion properties and genome The virions are spherical and 90–120 nm in diameter (Duarte et al. 1995; Mumford et al. 2003). The genome is composed of three RNAs (L RNA, 8955 nt, KM114546 = NC_027718; M RNA, 4830 nt, KM114547 = NC_027720; S RNA, 2947 nt, KM114548 = NC_027719) (Dullemans et al. 2015; Wu et al. 2015).

Chrysanthemum stunt viroid Taxonomic position Genus: Pospiviroid

(CSVd)

Family: Pospiviroidae

Geographical distribution CSVd infection in plants of Chrysanthemum spp. was reported in the early 1950s from the USA (Diener and Lawson 1973), and has spread to wherever chrysanthemum is grown (Horst et al. 1977; Lawson 1987; Dusi et al. 1990; Hill et al. 1996; Sugiura and Hanada 1998; Megan et al. 2001; Mathur et al. 2002; Chung et al. 2005; Matsushita et al. 2007; Mehle et al. 2010; Zhang et al. 2011b; Bostan et al. 2012; Cho et al. 2013; Zhao et al. 2015; Adkar-Purushothama et al. 2017; Van Bogaert et al. 2017). Symptoms and host(s) The viriod-infected chrysanthemum plants produce small and light green young leaves with chlorotic spots. Infected plants are reduced in size and they tend to flower earlier. Often the flowers are small and distorted and flower color is bleached. Susceptible cultivars often have numerous conspicuous white

Chrysanthemum spp.

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leaf spots, termed “measles.” CSVd has been found in mixed infection with some viruses (Hill et al. 1996; Hosokawa et al. 2004). The natural infections are reported in Chrysanthemum spp., Ageratum spp. (Z68201), Petunia hybrida, marguerite (Argyranthemum frutescens), Dahlia spp., Solanum jasminoides, Vinca spp., and eight wild Chrysanthemum spp. Transmission The viroid is transmitted by foliar contact, handling during cultivation (eg., disbudding) and through contaminated tools like cutting knives, scissors, etc. It is also transmitted mechanically through sap. The main hosts of CSVd are florists’ chrysanthemums (Dendranthema morifolium) and related ornamentals including Chrysanthemum praealtum, Dendranthema grandiflorum, D. indicum, and Tanacetum parthenium (Chung et al. 2001). Chung and Pak (2008) have reported seed transmission of this disease in chrysanthemum. The viroid could be disseminated over long distances in cuttings and other vegetative plants for planting. Etiology and genome properties The genome consists of a single-stranded circular RNA of 356 nt (V01107 = NC_002015) (Haseloff and Symons 1981; Gora-Sochacka 2004; Matsushita and Penmetcha 2009; Matsushita 2013; Yoon and Palukaitis 2013; Giguere et al. 2014b; Yoon et al. 2014).

Chrysanthemum vein chlorosis virus

(CVCV)

Synonyms Chrysanthemum frutescens virus (CFV) Taxonomic position CVCV is a tentative member of the family Rhabdoviridae. Geographical distribution CVSV infection in plants of Chrysanthemum spp. was reported from Brazil and Italy (Amici et al. 1978; Kitajima and Costa 1979). Symptoms and host(s) The virus-infected chrysanthemum plants exhibit symptoms of chlorotic banding of leaf veins. Transmission The virus could be disseminated over long distances in cuttings and other vegetative plants for planting. Virion properties and genome The virions are rhabdo or bullet shaped, with a clear modal length of 200–300 nm and 60–80 nm wide.

Chrysanthemum virus B Taxonomic position Genus: Carlavirus

(CVB)

Family: Betaflexiviridae

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504

Chrysanthemum spp.

Geographical distribution CVB is probably distributed worldwide (wherever Chrysanthemum spp. are grown) (Noordam 1952; Hollings 1957; Wilson and Moran 1983; Nakamura et al. 1994; Hill et al. 1996; Chung et al. 1999; Megan et al. 2001; Verma et al. 2003, 2007; Lin et al. 2005; Rishi 2009; Temaja et al. 2011; Zhao et al. 2015). Symptoms and host(s) The virus causes mild leaf mottling or vein-clearing in some chrysanthemum cultivars. In certain cases there is a loss of flower quality, and some cultivars, when infected with CVB, develop brown streaks on the florets of the compound flower. Mosaic, malformation, and slight-to-severe necrosis can also be found. Many chrysanthemum cultivars may be entirely infected with CVB without expressing symptoms (Hakkaart and Maat 1974). Transmission The virus is transmitted by aphid vectors Myzus persicae, Macrosiphum euphorbiae, Acyrthosiphon (Aulacorthum) solani, Coloradoa rufomaculata, and Macrosiphoniella (Pyrethromyzus) sanborni in a non-persistent manner (Temaja et al. 2011). The virus is transmissible by mechanical sap-inoculation, transmissible by grafting, not transmissible by contact between plants, and not transmissible by seed (Verma et al. 2003). The virus could be disseminated over long distances in cuttings and other vegetative plants for planting. Virion properties and genome The virions are flexuous filaments about 685 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8855 nt (NC_009087) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Levay and Zavriev 1991; Adams et al. 2004; Ohkawa et al. 2007; Singh et al. 2012).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV occurs wherever Chrysanthemum spp. plants are grown (Srivastava et al. 1992; Nakamura et al. 1994; Kumar et al. 2005; Verma et al. 2007; Rishi 2009; Kumar and Raj 2013; Zhao et al. 2015; Mitrofanova et al. 2018). The virus-infected chrysanthemum plants exhibit symptoms of diffused chlorosis, chlorotic dots near veins, and necrosis of leaves. At later stages of infection, older leaves show browning of major veins, leading to downward cupping and finally abscission. The virus is transmitted by aphid species such as Myzus persicae and Aphis gossypii and others in a non-persistent manner, and also by mechanical sap-inoculation to a wide host range. For more details of CMV, refer to Cucumis sativus.

Groundnut bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(GBNV)

Chrysanthemum spp.

505

GBNV infection in plants of Chrysanthemum indicum was reported from India (Holkar et al. 2013). The virus-infected chrysanthemum plants exhibit chlorotic patches, rings and necrosis of leaves, and bud and stem necrosis symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of GBNV, refer to Arachis hypogaea.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV occurs wherever Chrysanthemum spp. are grown (Louro 1996; Mertelik et al. 2002; Mavric and Ravnikar 2001; Shahraeen et al. 2002; Toth et al. 2007; Kondo et al. 2011; Toru et al. 2011; Ghotbi and Shahraeen 2012). The virus-infected chrysanthemum plants exhibit symptoms of chlorotic mottle, necrosis, distortion of young growth, lesions on the stem, and stunting and wilting of the plant. These symptoms can vary with the stage of growth of the host and with a variety of cultural conditions. The virus is mechanically sap-transmissible and by Western flower thrips Frankliniella occidentalis in a persistent-propagative manner. The virus could be disseminated over long distances in cuttings and other vegetative plants for planting. For more details of INSV, refer to Impatiens spp.

Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

IYSV was reported to infecting plants of Chrysanthemum spp. in Poland (Balukiewicz and Kryczynski 2005) and Iran (Rafizadeh et al. 2013). The virus-infected chrysanthemum plants exhibit chlorotic, necrotic, and diamond-shaped lesion symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of IYSV, refer to Iris spp. or Allium cepa.

Lettuce mosaic virus Taxonomic position Genus: Potyvirus

(LMV)

Family: Potyviridae

LMV was reported to infect Shasta daisy (Chrysanthemum maximum) in the USA (Koike and Davis 2009). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of LMV, refer to Lactuca sativa.

Pelargonium zonate spot virus Taxonomic position Genus: Anulavirus

(PZSV)

Family: Bromoviridae

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506

Chrysanthemum spp.

PZSV was reported to infect Chrysanthemum coronarium and Chrysanthemum segetum in Italy, causing foliar mosaic and bushy growth (Rana et al. 1990; Gallitelli et al. 2005). Thrips are involved in transmitting this virus by carrying the virus-infected pollen grains on their bodies and introducing the virus into the susceptible hosts while feeding. The virus is mechanically sap-transmissible to a wide range of test plants. For more details of PZSV, refer to Pelargonium spp.

Pepino mosaic virus Taxonomic position Genus: Potexvirus

(PepMV)

Family: Alphaflexiviridae

PepMV infection in plants of Chrysanthemum segetum was reported from Cyprus (Papayiannis et al. 2012). The virus is mechanically sap-transmissible. For more details of PepMV, refer to Solanum muricatum.

Potato virus Y

(PVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

PVY infection in plants of Chrysanthemum spp. was reported from China and Italy (Bertaccini et al. 1994; Liu et al. 2014; Zhao et al. 2015; Mitrofanova et al. 2018). The virus-infected chrysanthemum plants exhibit symptoms of leaf yellowing and mottled leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts and also by grafting. For more details of PVY, refer to Solanum tuberosum.

Soybean mosaic virus

(SMV)

Synonyms Chrysanthemum chlorotic spot virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

SMV infection in plants of Chrysanthemum frutescens was reported from Italy (Bertaccini et al. 1994). The virus causes marked chlorotic spots on leaves, but no symptoms on either stems or flowers, of Chrysanthemum frutescens, and was initially named as Chrysanthemum chlorotic spot virus (Bertaccini et al. 1994), but were subsequently identified as SMV (Bellardi et al. 1993). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SMV, refer to Glycine max.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

Chrysanthemum spp.

507

TMV infection in plants of Chrysanthemum spp. was reported from Egypt and China (Zhao et al. 2015; Nassar et al. 2012; Mitrofanova et al. 2018). There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Chrysanthemum morifolium and C. frutescens was reported from Iran (Ghotbi and Shahraeen 2009, 2012). The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Chrysanthemum spp. was reported from Iran (Ghotbi 2006; Vemana and Jain 2010). The virus-infected chrysanthemum plants exhibit mosaic, chlorosis, and leaf necrosis symptoms. The virus is transmitted by the thrips vectors. The virus present in/on pollen, entering into the host through the injuries caused by thrips while feeding. The virus is also mechanically sap-trasmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato aspermy virus

(TAV)

Synonyms Chrysanthemum aspermy virus; Chrysanthemum mild mottle virus Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

TAV is widespread and common in cultivated Chrysanthemum spp. (Mokra 1971; Horst et al. 1977; Chuyan and Krylov 1979; Gupta and Singh 1981; Wilson and Moran 1983; Raj et al. 1991, 2007, 2009; Hill et al. 1996; Chung et al. 1999; Megan et al. 2001; Lin et al. 2004; Verma et al. 2006, 2007, 2009; Kumar and Raj 2013; Zhao et al. 2015; Maddahian et al. 2017). The virus-infected chrysanthemum plants show symptoms of yellow mosaic, green vein-banding, chlorotic spots or ringspots and severe “flower break” or “color break” in flowers, distortion of flowers, and dwarfing of the chrysanthemum plants. Symptoms may not be expressed in the first year of infection, and most cultivars show no leaf symptoms or loss of plant vigor. These symptoms can vary with the stage of growth of the host and with a variety of cultural conditions. The virus is transmitted in a non-persistent manner through several aphid species such as Myzus persicae, Macrosiphoniella sanborni, Acyrthosiphon solani, and

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508

Chrysanthemum spp.

Aulacorthum solani. The virus is also mechanically sap-transmissible, and Chenopodium quinoa, Nicotiana glutinosa, N. tabacum, and Petunia spp. are suitable test plants. For more details of TAV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Chrysanthemum morifolium was reported from Iran (Ghotbi and Shahraeen 2009). The virus is transmitted by nematode vectors in a non-persistent manner. The virus is transmitted by mechanical sap-inoculation, and also by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV is present wherever chrysanthemum is cultivated (Wilson and Moran 1983; Matteoni and Allen 1989; Allen et al. 1990; Dal Bo et al. 1995; Hill et al. 1996; Roggero et al. 1998; Megan et al. 2001; Ravnikar et al. 2003; Balukiewicz et al. 2005; Morshedi Nour et al. 2012; Morshedi et al. 2013a, b; Stankovic et al. 2013; Marys et al. 2014; Renukadevi et al. 2015; El-Attar et al. 2017; Karavina and Gubba 2017). A number of strains of TSWV have been identified, and symptoms in chrysanthemum depend on the cultivar. Leaf symptoms range from ring and line patterns, faint mottling, and browning between leaf veins to yellowing of the veins. A characteristic necrosis starts in the leaf and extends through the petiole to the stem, forming longitudinal streaks. Flower distortion was also noticed (Matteoni and Allen 1989). The virus is spread from plant to plant by a thrips vector Frankliniella occidentalis in a persistent-propagative manner. The virus is acquired by the larvae when feeding on virus-infected plants, and is transmitted by both larval and adult thrips which remain infectious for life. The virus is mechanically sap-transmissible to many species of plants. Cuttings collected from infected plants are the main source of virus spread (Matsuura et al. 2004). For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Chrysanthemum spp. was reported from Cyprus (Papayiannis et al. 2011). The virus is transmitted by the whitefly vector, Bemisia tabaci in circulative non-propagative manner. The virus is transmitted by grafting but not transmitted by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

Chrysanthemum spp.

509

Tomato yellow ring virus

(TYRV)

Taxonomic position TYRV is a tentative member of the genus Orthotospovirus and family Tospoviridae TYRV infection in plants of Chrysanthemum spp. was reported from Iran (Ghotbi and Shahraeen 2012). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of TYRV, refer to Solanum lycopersicum.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of garland chrysanthemum (Chrysanthemum coronarium) was reported from Taiwan and Iran (Chen et al. 1999, 2000; Farzadfar et al. 2005). The virus-infected chrysanthemum plants show symptoms of albinism, mosaic, cleavage of leaves, and stunted growth. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TuMV, refer to Brassica rapa.

Watermelon bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(WBNV)

Family: Tospoviridae

WBNV infection in plants of Chrysanthemum indicum was reported from India (Holkar et al. 2017). The virus-infected chrysanthemum plants exhibit chlorotic patches, rings and necrosis of leaves, and bud and stem necrosis symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of WBNV, refer to Citrullus lanatus.

Zucchini yellow mosaic virus Taxonomic position Genus: Potyvirus

(ZYMV)

Family: Potyviridae

ZYMV infection in plants of Chrysanthemum morifolium was reported from China (Niu et al. 2015). The virus-infected chrysanthemum plants exhibit foliar chlorosis and mottle symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ZYMV, refer to Cucurbita pepo.

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Adkar-Purushothama CR, Chennappa G, Poornachandra Rao K, Sreenivasa MY, Nagendra Prasad MN, Maheshwar PK, Sano T (2015) Molecular identification of Chrysanthemum chlorotic mottle viroid infecting chrysanthemum in Karnataka, India. Plant Dis 99:1868 Adkar-Purushothama CR, Chennappa G, Poornachandra Rao K, Sreenivasa MY, Maheshwar PK, Nagendra Prasad MN, Sano T (2017) Molecular diversity among viroids infecting chrysanthemum in India. Virus Genes 53:636–642 Allen WR, Matteoni JA, Broadbent AB (1990) Susceptibility of cultivars of florist’s chrysanthemum to Tomato spotted wilt virus. Can J Plant Pathol 12:417–423 Amici A, Faoro F, Tornaghi R (1978) Natural occurrence of Rhabdovirus-like particles in thin sections of different plant species Avena sativa, Chrysanthemum frutescens, Holcus lanatus, Ranunculus repens, Secale cereale, oats, ornamental plant, rye. Riv Patol Veg 14:85–98 Balukiewicz A, Kryczynski S (2005) Tospoviruses in chrysanthemum mother stock plants in Poland. Phytopathol Pol 37:59–67 Balukiewicz A, Kryczynski S, Golnik K (2005) Biological differentation and molecular characterization of Tomato spotted wilt virus (TSWV) isolates from chrysanthemum plants. Phytopathol Pol 37:4–57 Bellardi MG, Marani F, Bertaccini A (1993) Detection of Soybean mosaic virus (SMV) in Chrysanthemum frutescens. Phytopathol Mediterr 32:156–158 Bertaccini A, Bellardi MG, Marani F, Rabiti A (1994) A Potyvirus infecting Chrysanthemum frutescens. Acta Hortic 377:107–114 Bezerra MI, Pozzer L, Nagata T, Lima MI, Katajima EW, de Avila AC, de Resende RO (1996) Chrysanthemum stem necrosis (CSNV), a proposed new species in the Tospovirus genus. Fitopatol Bras 21:430 Bezerra IC, de Resende RO, Pozzer L, Nagata T, Kormelink R, de Avila AC (1999) Increase of tospoviral diversity in Brazil with the identification of two new Tospovirus species, one from chrysanthemum and one from zucchini. Phytopathology 89:823–830 Bostan H, Gazel M, Elibuyuk IO (2012) Occurrence of pospiviroid in potato, tomato and some ornamental plants in Turkey. Afr J Biotechnol 9:2613–2617 CABI/EPPO (2005) Chrysanthemum stem necrosis virus, Distribution Maps of Plant Diseases, No. 947. CAB International, Wallingford Chen YK, Lee JY (2012) First report of Bidens mottle virus causing mosaic and leaf deformation in garland chrysanthemum and lettuce in Taiwan. Plant Dis 96:464 Chen TM, Chen YK, Chen MJ, Yeh SD (1999) Host reactions, cytological characteristics and serological properties of a Turnip mosaic virus isolate causing albinic mosaic disease of garland chrysanthemum. Plant Pathol Bull 8:73–82 Chen TM, Chen MJ, Yeh SD (2000) Characterization of the coat protein gene of a Turnip mosaic virus isolate infecting garland chrysanthemum. Plant Protect Bull Taipei 42:83–96 Cho WK, Jo Y, Jo K-M, Kim K-H (2013) A current overview of two viroids that infect chrysanthemums: Chrysanthemum stunt viroid and Chrysanthemum chlorotic mottle viroid. Viruses 5:1099–1113 Chung BN, Pak HS (2008) Seed transmission of Chrysanthemum stunt viroid in chrysanthemum (Dendranthema grandiflorum) in Korea. Plant Pathol J 23:334–338 Chung BN, Choi GS, Choi YM (1999) Identification of Tomato aspermy virus (TAV) and Chrysanthemum virus B (CVB) from Dendranthema indicum in Korea. Plant Pathol J 15(2):119–123 Chung BN, Choi GS, Kim HR, Kim JS (2001) Chrysanthemum stunt viroid in Dendranthema grandiflorum. Plant Pathol J 17:194–200 Chung BN, Lim JH, Choi SY, Kim JS, Lee EJ (2005) Occurrence of Chrysanthemum stunt viroid in chrysanthemum in Korea. Plant Pathol J 21:377–382 Chung BN, Kim DC, Kim JS, Cho JD (2006) Occurrence of Chrysanthemum chlorotic mottle viroid in chrysanthemum (Dendranthema grandiflorum) in Korea. Plant Pathol J 22(4):334–338 Chuyan AKH, Krylov AV (1979) Properties of Tomato aspermy virus from chrysanthemum and its host range in the Priomore region. Bull Clamogo Bot Kogosada 114:84–92 Dal Bo E, Ronco L, Alippi AM, Fernandez R (1995) Tomato spotted wilt virus on chrysanthemum in Argentina. Plant Dis 79:538 De Jonghe K, Morio S, Maes M (2013) First outbreak of Chrysanthemum stem necrosis virus (CSNV) on potted Chrysanthemum in Belgium. New Dis Rep 28:14 De la Pena M, Flores R (2002) Chrysanthemum chlorotic mottle viroid RNA: dissection of the pathogenicity determinant and comparative fitness of symptomatic and non-symptomatic variants. J Mol Biol 321:411–421 Di Serio F, Li S-F, Matousek J, Owens RA, Pallas V, Randles JW, Sano T, Verhoeven JThJ, Vidalakis G, Flores R, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Avsunviroidae. J Gen Virol 99:611–612 Diener TO, Lawson RH (1973) Chrysanthemum stunt: a viroid disease. Virology 51:94–101 Dimock AW, Geissinger CM, Horst RK (1971) Chlorotic mottle: a newly recognized disease of chrysanthemum. Phytopathology 61:415–419

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Duarte LML, Rivas EB, Alexandre MAV, de Avila AC, Nagata T, Chagas CM (1995) Chrysanthemum stem necrosis caused by a possible novel Tospovirus. J Phytopathol 143(9):569–571 Dullemans AM, Verhoeven JTJ, Kormelink R, van der Vlugt RAA (2015) The complete nucleotide sequence of Chrysanthemum stem necrosis virus. Arch Virol 160:605–608 Dusi AN, Fonseca MEN, Avila AC (1990) Occurrence of a viroid in chrysanthemum in Brazil. Plant Pathol 36:363–367 El-Attar AK, El-Shazly MA, Mokbel SA (2017) Characterization of a novel asymptomatic isolate of Tomato spotted wilt virus, infects chrysanthemum plants in Egypt. J Virol Sci 2:17–31 EPPO (2006) Chrysanthemum stem necrosis tospovirus, Datasheets on quarantine pests, web version 2006-03. EPPO, Paris 7 pp Farzadfar S, Ohshima K, Pourrahim R, Golnaraghi AR, Jalali S, Ahoonmanesh A (2005) Occurrence of Turnip mosaic virus on ornamental crops in Iran. Plant Pathol 54:261 Flores R, Daros JA, Hernandez C (2000) Avsunviroidae family: viroids containing hammerhead ribozymes. Adv Virus Res 55:271–323 Gallitelli D, Finetti-Sialer M, Martelli GP (2005) Anulavirus, a proposed new genus of plant viruses in the family Bromoviridae. Arch Virol 150:407–411 Ghotbi T (2006) First report on incidence of Tobacco streak virus (TSV) on ornamental plants in Iran. Iran J Plant Pathol 42:159–160 Ghotbi T, Shahraeen N (2005) First report on incidence of Arabis mosaic virus (ArMV, Nepovirus) on ornamental plants in Iran. Iran J Plant Pathol 41:305–306 Ghotbi T, Shahraeen N (2009) Natural incidence and infectivity level of three nepoviruses in ornamental crops in Iran. J Plant Breed Crop Sci 1:39–44 Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses affecting propagated ornamentals in Northern Iran. Int Res J Microbiol 3:373–381 Giguere T, Adkar-Purushothama CR, Bolduc F, Perreault JP (2014a) Elucidation of the structures of all members of the Avsunviroidae family. Mol Plant Pathol 15(8):767–779 Giguere T, Raj Adkar-Purushothama C, Perreault J-P (2014b) Comprehensive secondary structure elucidation of four genera of the family Pospiviroidae. PLoS One 9(6):e98655. https://doi.org/10.1371/journal.pone.0098655 Gora-Sochacka A (2004) Viroids: unusual small pathogenic RNAs. Acta Biochim Pol 51(3):587–607 Gupta RP, Singh BP (1981) Studies on Chrysanthemum aspermy virus on chrysanthemum. Indian J Mycol Pathol 11:158–160 Hakkaart FA, Maat FZ (1974) Variation of Chrysanthemum virus B. Neth J Plant Pathol 80:97–103 Haseloff J, Symons RH (1981) Chrysanthemum stunt viroid: primary sequence and secondary structure. Nucleic Acids Res 9:2741–2751 Hill MF, Giles RJ, Moran JR, Hepworth G (1996) The incidence of Chrysanthemum stunt viroid, Chrysanthemum B carlavirus, Tomato aspermy cucumovirus and Tomato spotted wilt tospovirus in Australian chrysanthemum crops. Aust Plant Pathol 25(3):174–178 Holkar SK, Mandal B, Jain RK (2013) New ornamental and cucurbitaceous hosts of Groundnut bud necrosis virus in India. Indian J Virol 24:146 Holkar SK, Kumar R, Yogita M, Katiyar A, Jain RK, Mandal B (2017) Diagnostic assays for two closely related Tospovirus species, Watermelon bud necrosis virus and Groundnut bud necrosis virus and identification of new natural hosts. J Plant Biochem Biotechnol 26:43–51 Hollings M (1957) Investigation of chrysanthemum viruses II. Virus B (mild mosaic) and Chrysanthemum Latent virus. Ann Appl Biol 45:589–602 Horst RK (1987) Chrysanthemum chlorotic mottle. In: Diener TO (ed) The viroids. Plenum, New York, pp 291–295 Horst RK, Langhans RW, Smith SH (1977) Effects of chrysanthemum stunt, chlorotic mottle, aspermy and mosaic on flowering and rooting of chrysanthemums. Phytopathology 67:9–14 Hosokawa M, Ueda E, Ohishi K, Otake A, Yazawa S (2004) Chrysanthemum stunt viroid disturbs the photoperiodic response for flowering of chrysanthemum plants. Planta 220:64–70 Karavina C, Gubba A (2017) Detection and characterization of Tomato spotted wilt virus infecting field and greenhousegrown crops in Zimbabwe. Eur J Plant Pathol 149:933–944 Kitajima EW, Costa AS (1979) Rhabdovirus-like particles in tissues of five different plant species. Fitopatol Bras 4:55–62 Koike ST, Davis RM (2009) UC IPM pest management guidelines: lettuce. University of California ANR Publication 3450, diseases. http://www.ipm.ucdavis.edu/PMG/r441101011.html. Accessed 12 May 2015 Kondo T, Yamashita K, Sugiyama S (2011) First report of Impatiens necrotic spot virus infecting chrysanthemum (Chrysanthemum morifolium) in Japan. J Gen Plant Pathol 77(4):263–265 Kumar S, Raj SK (2013) Molecular identification of cucumoviruses infecting chrysanthemums and their possible management. Indian J Virol 24:142 Kumar S, Srivastava A, Raj SK (2005) Molecular diagnosis of Cucumber mosaic virus in chrysanthemum. Indian J Virol 16:16

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Nakamura Y, Fujisawa I, Lee KH, Uematsu S (1994) Cucumber mosaic virus and Chrysanthemum virus B isolated from marguerite, Chrysanthemum frutescens L. Proc Kanto-Tosan Plant Protect Soc 41:177–179 Nassar EA, El-Dougdoug KA, Osman ME, Dawoud RA, Kinawy AH (2012) Characterization and elimination of a TMV isolate infecting chrysanthemum plants in Egypt. Int J Virol 8:14–26 Navarro B, Flores R (1997) Chrysanthemum chlorotic mottle viroid: unusual structural properties of a subgroup of selfcleaving viroids with hammerhead ribozymes. Proc Natl Acad Sci USA 94:11262–11267 Niu EB, Chen LJ, Niu YB (2015) First report of Zucchini yellow mosaic virus in chrysanthemum. Plant Dis 99:1289 Noordam D (1952) Virusziekten bij chrysant in nederland. With a summary: virus diseases of Chrysanthemum indicum in the Netherlands. Tijdschr Plantenziekten 58:121–190 Ohkawa A, Yamada M, Sayama H, Okuda S, Natsuaki T (2007) Complete nucleotide sequence of Japanese isolate of Chrysanthemum virus B (genus Carlavirus). Arch Virol 152:2253–2258 Okuda S, Okuda M, Matsuura S, Okazaki S, Iwai H (2013) Competence of Frankliniella occidentalis and Frankliniella intonsa strains as vectors for Chrysanthemum stem necrosis virus. Eur J Plant Pathol 136:355–362 Papayiannis L, Katis NI, Idris AM, Brown JK (2011) Identification of weed hosts of Tomato yellow leaf curl virus in Cyprus. Plant Dis 95:120–125 Papayiannis LC, Kokkinos CD, Alfafo-Fernandez A (2012) Detection, characterization and host range studies of Pepino mosaic virus in Cyprus. Eur J Plant Pathol 132:1–7 Rafizadeh N, Jafarpour B, Falahati Rastegar M (2013) Detection of Iris yellow spot virus (IYSV) in onion and some of ornamental plants by ELISA and RT-PCR methods in Khorasan Razavi provinces. 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ICPP, Christchurch Renukadevi P, Nagendran K, Nakkeeran S, Karthikeyan G, Jawaharlal M, Alice D, Malathi VG, Pappu HR (2015) First report of Tomato spotted wilt virus infection of chrysanthemum in India. Plant Dis 99:1190 Riley DG, Joseph SV, Srinivasan R, Diffie S (2011) Thrips vectors of tospoviruses. J Integr Pest Manag 2(1):1–10 Rishi N (2009) Significant plant virus diseases in India and a glimpse of modern disease management technology. J Gen Plant Pathol 75:1–18 Roggero P, Dellavalle G, Lisa V (1998) Tomato spotted wilt tospovirus in two cultivars of Chrysanthemum frutescens in Liguria. Colture Protette 27:67–68 Seo J-K, Kwak H-R, Kim M-K, Choi H-S (2015) First report of Carnation Italian ringspot virus in Chrysanthemum zawadskii var. latilobum in Korea. Plant Dis 99:1451 Shahraeen N, Ghotbi T, Mehraban AH (2002) Occurrence of Impatiens necrotic spot virus in ornamentals in Mahallat and Tehran provinces in Iran. Plant Dis 86:694 Singh BP, Gupta RP, Abidi SMH, Raizada RK (1978) Chlorotic mottle of Chrysanthemum morifolium (Ram.). A new record from India. Curr Sci 47:913 Singh L, Hallan V, Martin DP, Ram R, Zaidi AA (2012) Genomic sequence analysis of four new Chrysanthemum virus B isolates: evidence of RNA recombination. Arch Virol 157(3):531–537 Srivastava KM, Raj SK, Singh BP (1992) Properties of a Cucumber mosaic virus strain naturally infecting chrysanthemum in India. Plant Dis 76:474–477 Stankovic I, Bulajic A, Vucurovic A, Ristic D, Milojevic K, Nikolic D, Krstic B (2013) First report of Tomato spotted wilt virus on chrysanthemum in Serbia. Plant Dis 97:150 Sugiura H, Hanada K (1998) Chrysanthemum stunt viroid, a disease of large-flowered chrysanthemum (Dendranthema) in Niigata prefecture (Japan). 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Toth EK, Kriston E, Takacs A, Bajtek M, Kazinczi G, Horvath J (2007) First report of Impatiens necrotic spot virus in ornamental plants in Hungary. Plant Dis 91:331 Van Bogaert N, Smagghe G, Maes M, De Backer M, De Jonghe K (2017) Phylogeny of five predominant pospiviroid species in Belgium. Eur J Plant Pathol 149:25–33 Vemana K, Jain RK (2010) New experimental hosts of Tobacco streak virus and absence of true seed transmission in leguminous hosts. Indian J Virol 21:117–127 Verhoeven JTJ, Roenhorst JW, Cotes I, Peters D (1996) Detection of a novel tospovirus in chrysanthemum. Acta Hortic 432:44–51 Verma N, Sharma A, Ram R, Hallan V, Zaidi AA, Garg ID (2003) Detection, identification and incidence of Chrysanthemum B carlavirus in chrysanthemum in India. Crop Protect 22:425–429 Verma N, Hallan V, Ram R, Kumar K, Garg ID, Zaidi AA (2006) Detection and molecular characterization of a Tomato aspermy virus isolate infecting chrysanthemums in India. Acta Hortic 722:41–53 Verma N, Mehra A, Singh L, Hallan V, Singh AK, Jabeen N, Singh MK, Ram R, Zaidi AA (2007) Screening for viruses infecting chrysanthemum cultivars in India. Sci Hortic 111:260–265 Verma N, Kumar K, Kulshrestha S, Raikhy G, Hallan V, Ram R, Zaidi AA, Garg ID (2009) Molecular studies on Tomato aspermy virus isolates infecting chrysanthemums. Arch Phytopathol Plant Protect 42:99–111 Wilson JM, Moran JR (1983) The incidence of Chrysanthemum virus B, Tomato aspermy virus and Tomato spotted wilt virus in commercial chrysanthemum flower crops in Victoria. Aust Plant Pathol 12:17–19 Wu P-R, Chien W-C, Okuda M, Takeshita M, Yeh S-D, Wang Y-C, Chen T-C (2015) Genetic and serological characterization of Chrysanthemum stem necrosis virus, a member of the genus Tospovirus. Arch Virol 160:529–536 Yamamoto H, Sano T (2005) Occurrence of Chrysanthemum chlorotic mottle viroid in Japan. J Gen Plant Pathol 71:156–157 Yoon J-Y, Palukaitis P (2013) Sequence comparisons of global Chrysanthemum stunt viroid variants: multiple polymorphic positions scattered through the viroid genome. Virus Genes 46:97–104 Yoon J-Y, Cho I-S, Choi G-S, Choi S-K (2014) Construction of infectious cDNA clone of a Chrysanthemum stunt viroid Korean Isolate. Plant Pathol J 30:68–74 Yoon JY, Choi GS, Choi SK (2017) First report of Chrysanthemum stem necrosis virus on Chrysanthemum morifolium in Korea. Plant Dis 101:264 Zhang ZX, Pan S, Li SF (2011a) First report of Chrysanthemum chlorotic mottle viroid in chrysanthemum in China. Plant Dis 95:1320 Zhang Z-X, Ge B-B, Pan S, Zhao Z, Wang H-Q, Li S-F (2011b) Molecular detection and sequences analysis of Chrysanthemum stunt viroid. Acta Hortic Sin 38:2349–2356 Zhao X, Liu X, Ge B, Li M, Hong B (2015) A multiplex RT-PCR for simultaneous detection and identification of five viruses and two viroids infecting chrysanthemum. Arch Virol 160:1145–1152

Cicer arietinum (Chickpea) Family: Fabaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Grain legume

(AMV)

Family: Bromoviridae

AMV infection in plants of Cicer arietinum was reported from the USA, Africa, Asia, Australia, Iran, New Zealand, and Europe (Kaiser and Danesh 1971a, b; Fletcher 1987; Bos et al. 1988; Bosque-Perez and Buddenhagen 1990; Jones and Coutts 1996; Kawas et al. 1996; Latham and Jones 2001; Makkouk et al. 2001a, 2003a; Lava Kumar et al. 2008; van Leur et al. 2013). Early infection of chickpea plants results in the most severe symptoms, with shoot tip necrosis which extends throughout the plant resulting in death. With later infections, necrosis is usually restricted to the tip, and leaves lower down the infected shoot become chlorotic (pale) with reddening of leaf margins. Apical dominance

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breaks down, resulting in proliferation of axillary (side) shoots in some plants. Infected plants are stunted, shed their leaves, and die prematurely. In case of kabuli chickpea, the leaves develop chlorosis instead of reddening, and proliferation of axillary shoots is rarely observed. The virus is transmitted by three aphid species, Aphis craccivora, Acyrthosiphon pisum, and Acyrthosiphon sesbaniae, in a non-persistent manner (Kaiser and Danesh 1971a, b). The virus is also mechanically sap-transmissible. AMV is seed-borne in chickpea up to 0.1–1% (Jones and Coutts 1996). Seeds from diseased plants were often discolored, deformed, and shrivelled, and the germination was adversely affected. For more details of AMV, refer to Medicago sativa.

Bean leafroll virus

(BLRV)

Synonyms Pea leaf roll virus

Taxonomic position Genus: Luteovirus

Family: Luteoviridae

BLRV infection in plants of Cicer arietinum was reported from wherever the crop is grown (Kaiser and Danesh 1971a, b; Reddy et al. 1979; Bos et al. 1988; Brunt et al. 1990; Carazo et al. 1993; Nene et al. 1996; Makkouk et al. 2003a; Lava Kumar et al. 2008; Mustafayev et al. 2011). The virus-infected chickpea plants are easily spotted in the field by their yellow-, orange-, or brown-discolored foliage and stunted growth due to shortened internodes. In some cases stems show brown discoloration. Generally the tips and margins of leaflets are chlorotic before turning brown. The virus is transmitted by three aphid species, Aphis craccivora, Acyrthosiphon pisum, and A. sesbaniae, in a circulative, nonpropagative manner. The virus is not transmissible by mechanical sap-inoculation, and also by the seed. For more details of BLRV, refer to Phaseolus vulgaris.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Cicer arietinum was reported from the USA, Algeria, Iran, Puerto Rico, and India (Kaiser 1981; Chalam 1982; Chalam et al. 1985; Ait Yahia et al. 1997; Makkouk et al. 2003a; Singh et al. 2006). The virus-infected kabuli- and desi-type chickpeas develop apical necrosis, reddening of leaf margins, plant stunting, and premature senescence. Plants that are infected early but do not die may develop leaves that are filiform (having very narrow leaflets) and sometimes distorted. The virus is transmitted by three aphid species, Aphis craccivora, Acyrthosiphon pisum, and Acyrthosiphon sesbaniae, in a non-persistent manner, and which were primarily responsible for the spread of the virus (Kaiser and Danesh 1971a, b). The virus is mechanically sap-transmissible and infects a large number of hosts. BYMV in chickpea is not seed-borne; however, the seeds from diseased plants were often discolored, deformed, and shrivelled, and the germination was adversely affected (Kaiser and Danesh 1971a, b). For more details of BYMV, refer to Phaseolus vulgaris.

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Beet mosaic virus Taxonomic position Genus: Potyvirus

(BtMV)

Family: Potyviridae

BtMV was recorded in plants of Cicer arietinum from the Cap Bon region of Tunisia in 2006 (Kumari et al. 2010). The virus causes severe yellowing and tip wilting in chickpea. The legume aphid species Aphis fabae transmitted BtMV from chickpea to chickpea. The virus is mechanically sap-transmissible and also by grafting. The virus is not transmissible by contact between plants. For more details of BtMV, refer to Beta vulgaris.

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

BWYV infection in plants of Cicer arietinum was reported from Africa, Asia, Australia, Iran, Spain, Syria, Ethiopia, the USA, and Morocco (Bos et al. 1988; Bosque-Perez and Buddenhagen 1990; Carazo et al. 1993; Fortass et al. 1997; Makkouk et al. 1998, 2001a, 2002, 2003a; Tadesse et al. 1999a, b; Lava Kumar et al. 2008; Mustafayev et al. 2011; van Leur et al. 2013). The virus-infected chickpea plants exhibit yellowing and stunting with brown phloem discoloration. The virus is transmitted by aphid vectors, Myzus persicae, Aphis craccivora, and A. pisum, in a circulative and non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of BWYV, refer to Beta vulgaris.

Chickpea chlorosis Australia virus Taxonomic position Genus: Mastrevirus

(CpCAV)

Family: Geminiviridae

Geographical distribution CpCAV infection in plants of Cicer arietinum was reported from Australia (Schwinghamer et al. 2010; Thomas et al. 2010; Hadfield et al. 2012). Symptoms and host(s) The virus-infected chickpea plants exhibit stunting and leaf reddening or chlorosis symptoms. Transmission The insect vector of CpCAV has not been identified, but this is likely to be the leafhopper Orosius orientalis – the vector of other dicot-infecting mastreviruses in Australia. Transmission will likely be circulative and non-propagative. Similarly it is likely that the virus will prove not to be either mechanically or seed-transmitted. Virion properties and genome The structure of the virions of CpCAV has not been investigated. In common with all geminiviruses, the virions of CpCAV are likely geminate (twinned icosahedra).

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The genome of CpCAV consists of a single component of circular single-stranded DNA of ~2572 nt (JN989422 = NC_022131; GU256530) (Schwinghamer et al. 2010; Thomas et al. 2010). In common with all mastreviruses, CpYDV is predicted to encode four gene products, two in each orientation. Based on sequence homology to previously studied mastreviruses, these would code for the coat protein and movement protein in the virion-sense and in the complementary-sense for the replicationassociated protein (Rep) and Rep A (Palmer and Rybicki 1998; Boulton and Davies 2011; Hadfield et al. 2012; Muhire et al. 2013; Zerbini et al. 2017).

Chickpea chlorosis virus Taxonomic position Genus: Mastrevirus

(CpCV)

Family: Geminiviridae

Geographical distribution CpCV infection in plants of Cicer arietinum was reported from Australia (Schwinghamer et al. 2010; Thomas et al. 2010). Symptoms and host(s) The virus-infected chickpea plants exhibit stunting and leaf reddening or chlorosis symptoms. Transmission The insect vector of CpCV has not been identified, but this is likely to be the leafhopper Orosius orientalis – the vector of other dicot-infecting mastreviruses in Australia. Transmission will likely be circulative, non-propagative. Similarly, it is likely that the virus will prove not to be either mechanically or seed-transmitted. Virion properties and genome The structure of the virions of CpCV has not been investigated. In common with all geminiviruses the virions of CpCV will likely be geminated (twinned icosahedra). The genome of CpCV consists of a single component of circular single stranded DNA of ~2582 nt (GU256530 = NC_014740) (Schwinghamer et al. 2010; Thomas et al. 2010; Zerbini et al. 2017). In common with all mastreviruses, CpCV is predicted to encode four gene products, two in each orientation. Based on sequence homology to previously studied mastreviruses, these would code for the coat protein and movement protein in the virion-sense and in the complementary-sense for the replication-associated protein (Rep) and Rep A (Palmer and Rybicki 1998; Boulton and Davies 2011; Hadfield et al. 2012; Muhire et al. 2013).

Chickpea chlorotic dwarf virus Taxonomic position Genus: Mastrevirus

(CpCDV)

Family: Geminiviridae

Geographical distribution CpCDV was first reported on Cicer arietinum in India (Horn et al. 1993) and later in Egypt, Ethiopia, Iran, Iraq, Pakistan, Sudan, Syria, Tunisia, Africa, and Yemen (Horn et al. 1996; Hassan 2000; Hamed and Makkouk 2002; Makkouk et al. 1995, 2001a, 2002, 2003a; Kumari et al. 2004,

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2008, 2015; Lava Kumar et al. 2008). Various CpCDV isolates have previously been called Bean yellow darf virus, Chickpea chlorotic dwarf Syria virus and Chickpea chlorotic dwarf Sudan virus (Liu et al. 1997). Symptoms and host(s) The virus-infected chickpea plants cause leaf rolling, necrosis of the leaf tissue, yellowing or reddening of leaflets, phloem browning, and severe stunting. The virus is reported naturally infecting lentil, faba bean, tomato (Solanum lycopersicum), sugar beet (Beta vulgaris), bean (Phaseolus vulgaris), cucumber (Cucumis sativus), squash (Cucurbita pepo), and cotton (Gossypium hirsutum). Serologically the virus has also been identified infecting sugar beet (Beta vulgaris), bean (Phaseolus vulgaris), and faba bean (Vicia faba). Transmission The virus is transmitted by leafhopper vectors, Orosius orientalis and O. albicinctus, in a persistent, circulative, and non-propagative manner, and they acquire and transmit the virus within 2 h (Horn et al. 1993, 1994; Akhtar et al. 2011). The virus is not transmissible by mechanical sap-inoculation. Virion properties and genome Horn et al. (1993) purified CpCDV virions and showed these to have the typical geminivirus “geminated” (twinned quasi-icosahedral) morphology with a size of ~15  25 nm. Horn et al. (1993) showed the particles to contain a circular ssDNA molecule of. Since the work of Horn et al. (1993), the genomes of numerous isolates of CpCDV have been sequenced. The genome of CpCDV consists of a single component of circular single-stranded DNA of 2561 nt (Y11023 = NC_003493; AM933135, AM849097) (Horn et al. 1993; Palmer and Rybicki 1998; Hassan 2000; Nahid et al. 2008; Boulton and Davies 2011; Kraberger et al. 2013; Muhire et al. 2013; Zerbini et al. 2017). In common with all mastreviruses, CpCDV is predicted to encode four gene products, two in each orientation. Based on sequence homology to previously studied mastreviruses, these would code for the coat protein and movement protein in the virion-sense and in the complementary-sense for the replication-associated protein (Rep) and Rep A.

Chickpea chlorotic stunt virus Taxonomic position Genus: Polerovirus

(CpCSV)

Family: Luteoviridae

Geographical distribution CpCSV infection in plants of Cicer arietinum was reported from Tunisia, Azerbaijan, Ethiopia, Syria, Africa, and Iran (Abraham et al. 2006, 2009; Kumari et al. 2008; Asaad et al. 2009; Bananej et al. 2010; Mustafayev et al. 2011; Najar et al. 2011). Symptoms and host(s) The virus-infected chickpea plants exhibit yellowing, reddening, and stunting symptoms. Transmission The virus is transmitted by an aphid vector, Aphis craccivora, in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation.

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Virion properties and genome The virions are 25–30 nm in diameter, are hexagonal in outline, and have no envelope. The genome consists of single molecule of infectious, linear, positive-sense ssRNA of 5900 nt (NC_008249, AY956384) (Alnaasan et al. 2013). The 30 terminus has neither a poly(A) tract nor a tRNA-like structure. The 50 terminus has a genome-linked protein (VPg) (of about 17 kDa). The genome contains six open reading frames (ORFs).

Chickpea redleaf virus Taxonomic position Genus: Mastrevirus

(CpRLV)

Family: Geminiviridae

Geographical distribution CpRLV infection in plants of Cicer arietinum was reported from Australia (Schwinghamer et al. 2010; Thomas et al. 2010). Symptoms and host(s) The virus-infected chickpea plants exhibit stunting and leaf reddening or chlorosis symptoms. Transmission The insect vector of CpRLV has not been identified, but this is likely to be the leafhopper Orosius orientalis (syn. O. argentatus), the vector of other dicot-infecting mastreviruses in Australia. Transmission will likely be circulative and non-propagative. Similarly it is likely that the virus will prove not to be either mechanically or seed-transmitted. Virion properties and genome The structure of the virions of CpRLV has not been investigated. In common with all geminiviruses, the virions of CpRV are likely geminate (twinned icosahedra). The genome of CpRLV consists of a single component of circular single-stranded DNA of 2605 nt (GU256532 = NC_014739) (Palmer and Rybicki 1998; Thomas et al. 2010; Boulton and Davies 2011; Muhire et al. 2013; Zerbini et al. 2017). In common with all mastreviruses, CpRLV is predicted to encode four gene products, two in each orientation. Based on sequence homology to previously studied mastreviruses, these would code for the coat protein and movement protein in the virion-sense and in the complementary-sense for the replication-associated protein (Rep) and Rep A.

Chickpea stunt disease associated virus Taxonomic position Genus: Unassigned

(CpSDaV)

Family: Luteoviridae

Geographical distribution CpSDaV infection in plants of Cicer arietinum was reported from India (Naidu et al. 1997; Reddy and Kumar 2004; Lava Kumar et al. 2008). Symptoms and host(s) The virus-infected chickpea plants exhibit yellowing and stunting with brown phloem discoloration.

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Natural infection of this virus was reported only in chickpea. It is likely that in nature this virus may have several hosts; although since it causes symptomless infection, they may not be readily detectable. Transmission The virus is transmitted by aphid vectors, Aphis craccivora and Myzus persicae, in a circulative nonpropagative manner (Reddy and Kumar 2004). Virion properties and genome The virions are isometric with 28 nm diameter. The genome is single-stranded RNA (Reddy and Kumar 2004). A partial sequence is available (Y11530).

Chickpea yellow dwarf virus Taxonomic position Genus: Mastrevirus

(CpYDV)

Family: Geminiviridae

Geographical distribution CpYDV infection in plants of Cicer arietinum was reported from Pakistan (Kraberger et al. 2015). Symptoms and host(s) The virus has only been identified in chickpea exhibiting foliar yellowing symptoms. Transmission The vector of CpYDV has not been identified. It is likely that, in common with Chickpea chlorotic dwarf virus, the virus is transmitted by the leafhopper Orosius albicinctus. Virion properties and genome The structure of the virions of CpYDV has not been investigated. In common with all geminiviruses, the virions of CpYDV are likely geminate (twinned icosahedra). The genome of CpYDV consists of a single component of circular single-stranded DNA of ~2547 nt (KM377674 = NC_025475; KM377675) (Boulton and Davies 2011; Muhire et al. 2013; Kraberger et al. 2015; Zerbini et al. 2017). In common with all mastreviruses, CpYDV is predicted to encode four gene products, two in each orientation. Based on sequence homology to previously studied mastreviruses, these would code for the coat protein and movement protein in the virion-sense and in the complementary-sense for the replication-associated protein (Rep) and Rep A.

Chickpea yellow mosaic virus

(CpYMV)

Taxonomic position CpYMV is a tentative member of the genus Potyvirus and family Potyviridae Geographical distribution CpYMV infection in plants of Cicer arietinum was reported from Southern Bolivia (Larsen et al. 2003).

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Symptoms and host(s) The virus-infected chickpea plants exhibit symptoms consisting of mosaic, stunting, yellowing, wilting, shortening of internodes, and phloem discoloration. Transmission The virus is possibly transmitted by an aphid vector, Acyrthosiphon pisum. Virion properties and genome The virions are flexuous filaments. The genome consists of a single molecule of positive-sense singlestranded RNA. A partial genome sequence of 849 nt is available (AF527879) (Wylie et al. 2017).

Chickpea yellows virus Taxonomic position Genus: Mastrevirus

(CpYV)

Family: Geminiviridae

Geographical distribution CpYV infection in plants of Cicer arietinum was reported from Northeast Africa and Australia (Abraham et al. 2008; Hadfield et al. 2012). Symptoms and host(s) The symptoms exhibited by the plant were not described but are assumed to be a general chlorosis (yellowing) typical of chickpea-infecting mastreviruses. Transmission The insect vector of CpYV has not been identified, but this is likely to be the leafhopper Orosius orientalis (syn. O. argentatus) – the vector of other dicot-infecting mastreviruses in Australia. Transmission will likely be circulative and non-propagative. Similarly it is likely that the virus will prove not to be either mechanically or seed-transmitted. Virion properties and genome The structure of the virions of CpYV has not been investigated. In common with all geminiviruses, the virions of CpYV are likely geminate (twinned quasi-icosahedra). The genome of CpYV consists of a single component of circular single-stranded DNA of 2557 nt (JN989439) (Palmer and Rybicki 1998; Boulton and Davies 2011; Hadfield et al. 2012; Muhire et al. 2013; Zerbini et al. 2017). In common with all mastreviruses, CpYV is predicted to encode four gene products, two in each orientation. Based on sequence homology to previously studied mastreviruses, these would code for the coat protein and movement protein in the virion-sense and in the complementary-sense for the replication-associated protein (Rep) and Rep A.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

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CMV infection in plants of Cicer arietinum was first reported from Iran (Kaiser et al. 1968). The virus spreads in Iran, India, Pakistan, Morocco, Western Australia, Syria, and Spain (El-Maataoui and El-Hassani 1984; Chalam 1982; Chalam et al. 1986; Bashir and Malik 1993; Carazo et al. 1993; Jones and Coutts 1996; Latham and Jones 2001; Makkouk et al. 2001b, 2003a; Lava Kumar et al. 2008; van Leur et al. 2013). The virus-infected desi chickpea plants show initial symptoms of chlorosis of shoot tips, reduction in leaf size, and bunching of young leaves. Gradual reddening of the leaves then occurs with complete plant reddening in some varieties. Apical dominance breaks down, resulting in proliferation of axillary (side) shoots and stunting, with axillary shoots showing similar symptoms to those in the tip. The leaves of infected plants drop prematurely and the plants may die. In the case of kabuli chickpea, CMV infection exhibits distinct chlorosis in the leaves instead of reddening. Three aphid species, Aphis craccivora, Acyrthosiphon pisum, and Acyrthosiphon sesbaniae, were primarily responsible for the spread of virus in a non-persistent manner (Kaiser and Danesh 1971b). The virus is mechanically sap-transmissible and has a wide host range. CMV in chickpea is seed-borne up to 0.1–2% (Jones and Coutts 1996). Seeds from diseased plants were often discolored, deformed, and shrivelled, and the germination was adversely affected. For more details of CMV, refer to Cucumis sativus.

Faba bean necrotic yellows virus Taxonomic position Genus: Nanovirus

(FBNYV)

Family: Nanoviridae

FBNYV infection in plants of Cicer arietinum was reported from Asia and Africa (Horn et al. 1995; Makkouk et al. 1998, 2001a, 2002, 2003a, b; Kumari et al. 2008, 2009; Lava Kumar et al. 2008). The virus-infected chickpea plants exhibit symptoms of leaf rolling, yellowing (or reddening), and stunting. The virus is transmitted by aphid vectors, Acyrthosiphon pisum, Aphis craccivora, and A. fabae, in a persistent but non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of FBNYV, refer to Vicia faba.

Lettuce mosaic virus Taxonomic position Genus: Potyvirus

(LMV)

Family: Potyviridae

LMV infection in plants of Cicer arietinum was reported from California, USA (Bosque-Perez and Buddenhagen 1989). The virus-infected chickpea plants exhibit symptoms of mottling of leaves followed by tip wilting, general yellowing, and stem darkening symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of LMV, refer to Lactuca sativa.

Lettuce necrotic yellows cytorhabdovirus Taxonomic position Genus: Cytorhabdovirus

Family: Rhabdoviridae

(LNYV)

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LNYV infection in plants of Cicer arietinum was reported from Australia (Behncken 1983). The virusinfected chickpea plants exhibit symptoms of twisted shoots, and newly developed leaves may show a bleaching, necrotic tip burn, and/or leaflet chlorosis. The stem and bases of the larger petioles develop a reddish brown necrosis and the plants will wilt. The virus is transmitted by aphid vectors in a persistent, propagative manner, and also by mechanical sap-inoculation. For more details of LNYV, refer to Lactuca sativa.

Pea enation mosaic virus 1 Taxonomic position Genus: Enamovirus

(PEMV-1)

Family: Luteoviridae

PEMV-1 infection in plants of Cicer arietinum was recorded in Syria, Puerto Rico, Iran, the USA, and Italy (Kaiser 1981; Makkouk et al. 2001a, 2003a). Chickpea plants infected with PEMV-1 typically are severely stunted showing twisted and malformed leaves. Pods also are misshapen, are poorly filled, and often show outgrowths that appear as bumps on the surface. Plants infected at an early growth stage may die. Infected plants rarely yield seed of any quality. The virus is transmitted by aphid vectors Acyrthosiphon pisum and Myzus persicae in a persistent manner, and also by mechanical sap-inoculation. For more details of PEMV-1, refer to Pisum sativum.

Pea seed-borne mosaic virus Taxonomic position Genus: Potyvirus

(PSbMV)

Family: Potyviridae

PSbMV infection in plants of Cicer arietinum was reported from Morocco, Pakistan, Iran, and Ethiopia (Ouizbouben and Fortass 1997; Tadesse et al. 1999b; Makkouk et al. 2001b, 2003a). The virus-infected chickpea plants show symptoms of very narrow (filiform) leaflets that may be twisted or turned down, indistinct mosaic, mottling, necrotic lesions on leaflets, necrosis of shoot tips, stunting of plants, proliferation of stems, and abortion of podding. The severity of symptoms varies with isolate and stage of plants infected. The virus is transmitted by a number of aphid species in a non-persistent manner. It is also transmissible by mechanical sap-inoculation, and seed transmission is also reported (Makkouk et al. 2001b). For more details of PSbMV, refer to Pisum sativum.

Pea streak virus Taxonomic position Genus: Carlavirus

(PeSV)

Family: Betaflexiviridae

PeSV was first detected in plants of Cicer arietinum in the Palouse region of Eastern Washington State (USA) in 1979 (Bos et al. 1988; Kaiser et al. 1993; Larsen et al. 1993). The virus-infected chickpea plants exhibit symptoms of yellowing of the foliage, stunting, necrosis of leaflets, wilting of terminal shoots, and phloem discoloration. Infected seedlings wilt and die prior to pod formation, and the seeds in infected mature plants are small, discolored, and misshapen. The virus is transmitted by aphid vectors

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in a non-persistent manner, and also by mechanical sap-inoculation; no seed or pollen transmission is reported. For more details of PeSV, refer to Pisum sativum.

Red clover vein mosaic virus Taxonomic position Genus: Carlavirus

(RCVMV)

Family: Betaflexiviridae

RCVMV infection in plants of Cicer arietinum was reported from Washington State, USA (Larsen et al. 1996; Larsen and Miklas 2001). The virus-infected chickpea plants exhibit symptoms of severe stunting, mosaic, proliferation of axillary buds, and malformation of leaves and branches. Flower and pod formation numbers on affected plants were severely reduced. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a number of hosts. For more details of RCVMV, refer to Trifolium spp.

Soybean dwarf virus

(SbDV)

Taxonomic position Genus: Luteovirus

Family: Luteoviridae

SbDV infection in plants of Cicer arietinum was reported from Iran and Ethiopia (Bos et al. 1988; Tadesse et al. 1999a; Makkouk et al. 2002, 2003a). The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation, and also not by contact between plants. For more details of SbDV, refer to Glycine max.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Cicer arietinum was reported from India (Dhingra 1981). The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Cicer arietinum was reported from Australia (Sharman et al. 2008, 2015). The virus-infected chickpea plants exhibit symptoms of stem necrosis, tip wilting, and necrotic etching on the leaves. The virus is transmitted by the thrips vectors. The virus present in/on pollen, and enters into the host through the injuries caused by thrips while feeding. The virus is mechanically sap-

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transmissible and is not transmissible by contact between plants. The virus is transmissible through the seed in chickpea up to 11.1% (Kaiser et al. 1991). For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Cicer arietinum was reported from Brazil and Australia (Boiteux et al. 1995; Thomas et al. 2004). The virus-infected chickpea plants exhibit symptoms of wilting and/or necrosis of shoot tips or chlorosis and bunching of shoot tips accompanied by reddening, chlorosis, and subsequent death of entire plants. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Cicer arietinum was reported from Australia (Schwinghamer et al. 2007; van Leur et al. 2013). The virus-infected chickpea plants exhibit symptoms of wilting and/or necrosis of shoot tips, concomitant with or leading to reddening, chlorosis, and premature death of entire plants. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

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Kumari SG, Makkouk KM, Loh MH, Negassi K, Tsegay S, Kidane R, Kibert A, Tesfatsion Y (2008) Virus diseases affecting chickpea crops in Eritrea. Phytopathol Mediterr 47:42–49 Kumari SG, Attar N, Mustafayev E, Akparov Z (2009) First report of Faba bean necrotic yellows virus affecting legume crops in Azerbaijan. Aust Plant Dis Notes 93:1220 Kumari SG, Najar A, Attar N, Loh MH, Vetten H-J (2010) First report of Beet mosaic virus infecting chickpea (Cicer arietinum) in Tunisia. Plant Dis 94:1068 Kumari SG, Najar A, Timoumi S, Male MF, Kraberger S, Varsani A (2015) First report of chickpea chlorotic dwarf virus naturally infecting chickpea in Tunisia. New Dis Rep 32:16 Larsen RC, Miklas PN (2001) Effect of Red clover vein mosaic carlavirus infection on seed production and biomass yield in chickpea. Phytopathology 91:S186 Larsen RC, Wyatt SD, Kaiser WJ (1993) Characterization of a severe strain of Pea streak carlavirus isolated from chickpea in Washington. Phytopathology 83:1374 Larsen RC, Kaiser WJ, Wyatt SD (1996) First report of a virus disease of chickpea caused by a strain of Red clover vein mosaic carlavirus. Plant Dis 80:709 Larsen RC, Kaiser WJ, Wyatt SD, Buxton-Druffel KL, Berger PH (2003) Characterization of a new potyvirus naturally infecting chickpea. Plant Dis 87:1366–1371 Latham LJ, Jones RAC (2001) Incidence of virus infection in experimental plots, commercial crops and seed stocks of cool season crop legumes. Aust J Agric Res 52:397–413 Lava Kumar P, Kumari SMG, Waliyar F (2008) Virus diseases of chickpea. In: Rao GP, Kumar PL, Holguin-Pena RJ (eds) Characterization, diagnosis and management of plant viruses. Studium Press, pp 213–234 Liu L, van Tonder T, Pietersen G, Davies JW, Stanley J (1997) Molecular characterization of a subgroup I geminivirus from a legume in South Africa. J Gen Virol 78(PT 8):2113–2117 Makkouk KM, Dafalla G, Hussein M, Kumari SG (1995) The natural occurrence of Chickpea chlorotic dwarf geminivirus in chickpea and faba bean in the Sudan. J Phytopathol 143:465–466 Makkouk KM, Bashir M, Jones R (1998) First record of Faba bean necrotic yellows virus and Beet western yellows luteovirus affecting lentil and chickpea in Pakistan. Plant Dis 82:591 Makkouk KM, Bashir M, Jones RAC, Kumari SG (2001a) Survey for viruses in lentil and chickpea crops in Pakistan. Z Pflanzenkr Pflanzenschutz 108:258–268 Makkouk KM, Kumari SG, Lesemann DE (2001b) First record of Pea enation mosaic virus naturally infecting chickpea and grasspea crops in Syria. Plant Dis 85:1032 Makkouk KM, Fazlali Y, Kumari SG, Farzadfar S (2002) First record of Beet western yellows virus, Chickpea chlorotic dwarf virus, Faba bean necrotic yellows virus and Soybean dwarf virus infecting chickpea and lentil crops in Iran. Plant Pathol 51:387 Makkouk KM, Hamed AA, Hussein M, Kumari SG (2003a) First report of Faba bean necrotic yellows virus (FBNYV) infecting chickpea (Cicer arietinum) and faba bean (Vicia faba) crops in Sudan. Plant Pathol 52(3):412 Makkouk KM, Kumari SG, Shahreen N, Fazlali Y, Farzadfar S, Ghotbi T, Reza Mansouri A (2003b) Identification and seasonal variation of viral diseases of chickpea and lentil in Iran. J Plant Dis Protect 110:157–169 Muhire B, Martin DP, Brown JK, Navas-Castillo J, Moriones E, Zerbini FM, Rivera-Bustamante R, Malathi VG, Briddon RW, Varsani A (2013) A genome-wide pairwise-identity-based proposal for the classification of viruses in the genus mastrevirus (family Geminiviridae). Arch Virol 158(6):1411–1424 Mustafayev E, Kumari SG, Attar N, Zeynal A (2011) Viruses infecting chickpea and lentil crops in Azerbaijan. Australas Plant Pathol 40(6):612–620 Nahid N, Amin I, Mansoor S, Rybicki EP, van der Walt E, Briddon RW (2008) Two dicot-infecting mastreviruses (family Geminiviridae) occur in Pakistan. Arch Virol 153(8):1441–1451 Naidu RA, Mayo MA, Reddy SV, Jolly CA, Torrence L (1997) Diversity among coat proteins of luteoviruses associated with chickpea stunt disease in India. Ann Appl Biol 130:37–47 Najar A, Kumari S, Attar N, Lababidi S (2011) First report of Chickpea chlorotic stunt virus infecting legume crops in Tunisia. Plant Dis 95:1321 Nene YL, Sheila VK, Sharma SB (1996) A world list of chickpea (Cicer arietinum) and pigeonpea (Cajanus cajan). Pathogens, 5th edn. ICRISAT Publication, Hyderabad, India, 25 pp Ouizbouben A, Fortass M (1997) Survey of chickpea for viruses in Morocco. EPPO Bull 27:249–254 Palmer KE, Rybicki EP (1998) The molecular biology of mastreviruses. Adv Virus Res 50:183–234 Reddy SV, Kumar PL (2004) Transmission and properties of a new luteovirus associated with chickpea stunt disease in India. Curr Sci 86:1157–1161 Reddy MV, Nene YL, Verma JP (1979) Pea leaf roll causes chickpea stunt. Int Chickpea Newsl 1:8 Schwinghamer MW, Thomas JE, Parry JN, Schilg MA, Dann EK (2007) First record of natural infection of chickpea by Turnip mosaic virus. Aust Plant Dis Notes 2:41–43

C

528

Cichorium endivia (Endive)

Schwinghamer MW, Thomas JE, Schilg MA, Parry JN, Dann EK, Moore KJ, Kumari SG (2010) Mastreviruses in chickpea (Cicer arietinum) and other dicotyledonous crops and weeds in Queensland and northern New South Wales, Australia. Aust Plant Pathol 39:551–561 Sharman M, Thomos JE, Persley DM (2008) First report of Tobacco streak virus in sunflower (Helianthus annuus), cotton (Gossypium hirsutum), chickpea (Cicer arietinum), mungbean (Vigna radiata) in Australia. Aust Plant Dis 3:27–29 Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct Tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207 Singh A, Rishi N, Sirohi A (2006) A narrow leaf disease of chickpea in Himachal Pradesh caused by a potyvirus. Indian J Crop Sci 1:178–179 Tadesse N, Ali K, Gorfu D, Abraham A, Lencho A, Ayalew M, Yusuf A, Makkouk KM, Kumari SG (1999a) First report of Soybean dwarf virus infecting lentil and beet western yellows virus infecting lentil and chickpea crops in Ethiopia. Plant Dis 83:589 Tadesse N, Ali K, Gorfu D, Yusuf A, Abraham A, Ayalew M, Lencho A, Makkouk KM, Kumari SG (1999b) Survey for chickpea and lentil virus diseases in Ethiopia. Phytopathol Mediterr 38:149–158 Thomas JE, Schwinghamer MW, Parry JN, Sharman M, Schilg MA, Dann EK (2004) First report of Tomato spotted wilt virus in chickpea (Cicer arietinum) in Australia. Australas Plant Pathol 33:597–599 Thomas JE, Parry JN, Schwinghamer MW, Dann EK (2010) Two novel mastreviruses from chickpea (Cicer arietinum) in Australia. Arch Virol 155:1777–1788 van Leur JAG, Aftab M, Manning W, Bowring A, Riley MJ (2013) A severe outbreak of chickpea viruses in northern New South Wales, Australia, during 2012. Australas Plant Dis Notes 8:49–53 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Cichorium endivia (Endive) Family: Asteraceae

Bidens mottle virus Taxonomic position Genus: Potyvirus

Leafy vegetable

(BiMoV)

Family: Potyviridae

BiMoV infection in plants of Cichorium endivia was reported from Florida, USA (Purcifull et al. 1971). The virus-infected endive plants exhibit symptoms of mottled leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BiMoV, refer to Bidens spp.

Endive necrotic mosaic virus Taxonomic position Genus: Potyvirus

(ENMV)

Family: Potyviridae

Geographical distribution ENMV was first reported in plants of Cichorium endivia from Germany by Vetten et al. (1995). The virus spreads in Germany. Symptoms and host(s) The virus-infected endive plants exhibit severe mosaic and necrosis symptoms.

Cichorium endivia (Endive)

529

Transmission The virus is transmitted by an aphid vector, Myzus persicae, in a non-persistent manner. The virus is also transmissible by mechanical sap-inoculation. Virion properties and genome The virions are non-enveloped, flexuous filaments, with a clear modal length of 820 nm and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of c.9.5–10 kb (AJ223827) (Revers and Garcia 2015). A partial genome sequence of 1346 nt is available (AJ223827) (Wylie et al. 2017).

Lettuce big-vein associated varicosavirus Taxonomic position Genus: Varicosavirus

(LBVaV)

Family: Rhabdoviridae

LBVaV infection in plants of Cichorium endivia was reported from Brazil (Colariccio et al. 2005). The virus-infected endive plants exhibit symptoms of chlorotic thickening of foliar veins, defective growth, and, in some cases, failure to form complete heads. The virus is transmitted by a fungal vector Olpidium brassicae and also by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants. For more details of LBVaV, refer to Lactuca sativa.

Mirafiori lettuce big-vein ophiovirus Taxonomic position Genus: Ophiovirus

(MiLBVV)

Family: Aspiviridae

MiLBVV infection in plants of Cichorium endivia was reported from Brazil (Colariccio et al. 2005). The virus-infected endive plants exhibit symptoms of chlorotic thickening of foliar veins, defective growth, and, in some cases, failure to form complete heads. The virus is transmitted by a fungal vector Olpidium brassicae and also by mechanical sap-inoculation. For more details of MiLBVV, refer to Lactuca sativa.

Plum pox virus Taxonomic position Genus: Potyvirus

(PPV)

Family: Potyviridae

PPV infection in plants of Cichorium endivia was reported from Slovenia (Virscek Marn et al. 2004). The virus-infected endive plants exhibit pale or yellow green ringspots or mottling on the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of PPV, refer to Prunus domestica.

Tomato chlorotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(TCSV)

C

530

Cichorium endivia ssp. latifolium (Escarole)

TCSV infection in plants of Cichorium endivia was reported from Brazil (Colariccio et al. 2001). The virus-infected endive plants exhibit symptoms of mosaic, necrosis, chlorotic and necrotic ringspots, and a reduction in leaf size. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of TCSV, refer to Solanum lycopersicum.

References Colariccio A, Chaves ALR, Eiras M, Chagas CM (2001) Identification of the Tomato chlorotic spot virus in endive (Cichorium endivia L.). Summa Phytopathol 27:325–327 Colariccio A, Chaves ALR, Eiras M, Chagas CM, Roggero P (2005) Detection of Varicosavirus and Ophiovirus in lettuce associated with lettuce big-vein symptoms in Brazil. Fitopatol Bras 30:416–419 Purcifull DE, Christie SR, Zitter TA, Bassett MJ (1971) Natural infection of lettuce and endive by Bidens mottle virus. Plant Dis Reptr 55:1061–1063 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Vetten HJ, Lesemann DE, Schonfelder M (1995) Proceedings of the 8th conference on virus diseases of vegetables, Prague, 9–15 July 1995, p 113 Virscek Marn M, Mavric I, Urbancic-Zemljic M, Skerlavaj V (2004) Detection of Plum pox virus Potyvirus in weeds. Acta Hortic 657:251–254 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Cichorium endivia ssp. latifolium (Escarole) Family: Asteraceae

Leafy vegetable

Lily mottle virus Taxonomic position Genus: Potyvirus

(LMoV)

Family: Potyviridae

LMoV infection in plants of Cichorium endivia ssp. latifolium was reported from Italy (Lisa et al. 2002). The virus-infected escarole plants exhibit symptoms of mosaic and necrotic spots on outer leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of LMoV, refer to Lilium spp.

Moroccan pepper virus Taxonomic position Genus: Tombusvirus

(MPV)

Family: Tombusviridae

MPV infection in plants of Cichorium endivia ssp. latifolium plants was reported from New Jersey (USA) by Wintermantel and Bachinsky (2014). The virus-infected escarole plants had cores of the heads which were necrotic and rotted, while normally white outer leaves which were yellow with more pronounced yellowing of the veins and occasional veinal necrosis. The virus is transmissible by

Cichorium intybus (Chicory)

531

mechanical sap-inoculation to three to nine families. The virus is transmissible by grafting. There is no known vector for this virus. For more details of MPV, refer to Capsicum annuum.

Plum pox virus Taxonomic position Genus: Potyvirus

(PPV)

C Family: Potyviridae

PPV infection in plants of Cichorium endivia ssp. latifolium was reported from Slovenia (Virscek Marn et al. 2004). The virus-infected escarole plants exhibit pale or yellow green ringspots or mottling on the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PPV, refer to Prunus domestica.

Tomato infectious chlorosis virus Taxonomic position Genus: Crinivirus

(TICV)

Family: Closteroviridae

TICV infection in plants of Cichorium endivia ssp. latifolium was reported from Italy (Parrella and Filella 2007; Parrella 2008). The virus-infected escarole plants exhibit symptoms of interveinal yellowing of the leaves. The virus is transmitted by the whitefly vector, Trialeurodes vaporariorum, in a semi-persistent manner. The virus is not transmitted by mechanical sap-inoculation. For more details of TICV, refer to Solanum lycopersicum.

References Lisa V, Vetten HJ, Lesemann D-E, Gotta P (2002) Occurrence of Lily mottle virus in Escarole. Plant Dis 86:329 Parrella G (2008) Interveinal yellowing caused by Tomato infectious chlorosis virus in lettuce and escarole in southern Italy. J Phytopathol 156:190–192 Parrella G, Filella F (2007) A severe outbreak of Tomato infectious chlorosis virus in lettuce and escarole in Southern Italy. J Plant Pathol 89(3):S52 Virscek Marn M, Mavric I, Urbancic-Zemljic M, Skerlavaj V (2004) Detection of Plum pox virus potyvirus in weeds. Acta Hortic 657:251–254 Wintermantel WM, Bachinsky D (2014) First report of Moroccan pepper virus in association with yellows on escarole in the United States and the world. Plant Dis 98:1448

Cichorium intybus (Chicory) Family: Asteraceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Medicinal

(AMV)

Family: Bromoviridae

532

Cichorium intybus (Chicory)

AMV infection in plants of Cichorium intybus was reported from Saudi Arabia (Al-Shahwan et al. 2017). The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Artichoke Italian latent virus Taxonomic position Genus: Nepovirus

(AILV)

Family: Secoviridae

AILV infection in plants of Cichorium intybus was reported from Apulia (Italy) by Vovlas et al. (1971). The virus-infected chicory plants show chlorotic mottling of the leaves often with bright yellow spots. The virus is transmitted by nematode vectors, Longidorus fasciatus and L. apulus, in a non-persistent manner, and also by mechanical sap-inoculation (Brown and Trudgill 1998). For more details of AILV, refer to Cynara cardunculus var. scolymus.

Chicory yellow mottle virus Taxonomic position Genus: Nepovirus

(ChYMV)

Family: Secoviridae

Geographical distribution ChYMV was first reported in plants of Cichorium intybus from Italy by Vovlas et al. (1971). The virus spreads in Italy (Quacquarelli et al. 1972; Piazzolla et al. 1986). Symptoms and host(s) The virus-infected chicory plants exhibit symptoms of ringspots, line pattern, and bright yellow mottling of leaves. Transmission The virus is possibly transmitted by nematode vectors in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are isometric and non-enveloped of two types but similar in size 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is bipartite, linear, positive-sense, single-stranded RNA. The complete genome sequence is not available; however two satellite RNAs associated with ChYMV were characterized (Rubino et al. 1990). The large satellite RNA is 1145–1165 nt (D00686 = NC_003778; D00722) and has a 50 leader sequence, a long open reading frame coding for a protein of 39.6 kDa, a 30 noncoding region, and a poly(A) tail. The small satellite RNA (linear and circular forms) is 457 nt (D00685 = NC_003971, D00721) and has no significant coding region (Quacquarelli et al. 1972; Rubino et al. 1990; Sanfacon et al. 2009; Dunez and Le Gall 2011; Sanfacon 2015; Thompson et al. 2017).

Cichorium intybus (Chicory)

Plum pox virus Taxonomic position Genus: Potyvirus

533

(PPV)

Family: Potyviridae

PPV infection in plants of Cichorium intybus was reported from Slovenia (Virscek Marn et al. 2004). The virus-infected chicory plants exhibit pale or yellow green ringspots or mottling on the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of PPV, refer to Prunus domestica.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Cichorium intybus was reported from Italy (Grieco et al. 2000). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sapinoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Al-Shahwan IM, Abdalla OA, Al-Saleh MA, Amer MA (2017) Detection of new viruses in alfalfa, weeds and cultivated plants growing adjacent to alfalfa fields in Saudi Arabia. Saudi J Biol Sci 24(6):1336–1343 Brown DJF, Trudgill DL (1998) Nematode transmission of plant viruses: a 30 year perspective. Host pathogen interactions and crop protection. SCRI Annual Report. pp 121–125 Dunez J, Le Gall O (2011) Nepovirus. Comoviridae. In: The Springer index of Viruses. Springer, New York, pp 361–369. https://doi.org/10.1007/978-0-387-95919-1_55 Grieco PD, Conte D, Munno I, Nuzzaci M, de Stradis A (2000) Tomato spotted wilt virus (TSWV) on weeds and wild plants in Metapontino, Basilicata. Informatore Fitopatol 50:43–46 Piazzolla P, Vovlas C, Rubino L (1986) Symptom regulation induced by Chicory yellow mottle virus satellite-like RNA. J Phytopathol 115:124–129 Quacquarelli A, Vovlas C, Piazzolla P, Russo M, Martelli GP (1972) Some characteristics of Chicory yellow mottle virus. Phytopathol Mediterr 11(3):180–188 Rubino L, Tousignant ME, Steger G, Kaper JM (1990) Nucleotide sequence and structural analysis of two satellite RNAs associated with Chicory yellow mottle virus. J Gen Virol 71:1897–1903 Sanfacon H (2015) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. In: eLS. Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000764.pub3. http://www.els.net Sanfacon H, Wellink J, Le Gall O, Karasev A, van der Vlugt R, Wetzel T (2009) Secoviridae: a proposed family of plant viruses within the order Picornavirales that combines the families Sequiviridae and Comoviridae, the unassigned genera Cheravirus and Sadwavirus, and the proposed genus Torradovirus. Arch Virol 154(5):899–907 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J Gen Virol 98:529–531 Virscek Marn M, Mavric I, Urbancic-Zemljic M, Skerlavaj V (2004) Detection of Plum pox virus Potyvirus in weeds. Acta Hortic 657:251–254 Vovlas C, Martelli GP, Quacquarelli A (1971) Le virosi delle piante ortensi in Puglia. VI. II complesso delle maculature anulari della cicoria. Phytopathol Mediterr 10:244–254

C

534

Cirsium arvense (Creeping thistle)

Cirsium arvense (Creeping thistle) Family: Asteraceae

Weed host

Alfalfa mosaic virus

(AMV)

Taxonomic position Genus: Alfamovirus

Family: Bromoviridae

AMV was reported from New Zealand infecting Cirsium arvense plants without obvious virus symptoms (Fletcher 2001). The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Amaranthus leaf mottle virus Taxonomic position Genus: Potyvirus

(AmLMV)

Family: Potyviridae

AmLMV infection in plants of Cirsium arvense was reported by Casetta et al. (1986). The virusinfected creeping thistle plants exhibit leaf mottling symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AmLMV, refer to Amaranthus spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Cirsium arvense was reported from Italy and Bulgaria (Vicchi and Bellardi 1988; Dikova 1989). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Plum pox virus Taxonomic position Genus: Potyvirus

(PPV)

Family: Potyviridae

PPV infection in plants of Cirsium arvense was reported from Slovenia (Virscek Marn et al., 2004; Milusheva and Rankova 2006). The virus-infected creeping thistle plants exhibit pale or yellow green ringspots or mottling on the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PPV, refer to Prunus domestica.

Cirsium arvense (Creeping thistle)

Thistle mottle virus Taxonomic position Genus: Caulimovirus

535

(ThMoV)

Family: Caulimoviridae

Geographical distribution ThMoV infection in plants of Cirsium arvense was first reported from Norwich, UK, by Donson and Hull (1983). Symptoms and host(s) The virus-infected creeping thistle plants exhibit slight mottling symptoms. Transmission The virus is transmitted by aphid vectors in a semi-persistent manner, and also through mechanical sapinoculation. Virion properties and genome The virions are isometric, non-enveloped, and 45–50 nm in diameter with no obvious surface structure. The genome is a monopartite, circular, double-stranded DNA of 7.8 kb (Hohn 2011).

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Cirsium arvense was reported from Italy (Grieco et al. 2000). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Casetta AD, Agostino G, Conti M (1986) Isolation of Amaranthus leaf mottle virus (ALMV) from Cirsium arvense Scop. Informatore Fitopatol 36(6):43–46 Dikova B (1989) Wild-growing hosts of the Cucumber mosaic virus (abstract). Rasteniev’dni Nauki 26(7):57–64 Donson J, Hull R (1983) Physical mapping and molecular cloning of Caulimovirus DNA. J Gen Virol 64:2281–2288 Fletcher D (2001) New hosts of Alfalfa mosaic virus, Cucumber mosaic virus, Potato virus Y, Soybean dwarf virus, and Tomato spotted wilt virus in New Zealand. N Z J Crop Hortic Sci 29:213–217 Grieco PD, Conte D, Munno I, Nuzzaci M, de Stradis A (2000) Tomato spotted wilt virus (TSWV) on weeds and wild plants in Metapontino, Basilicata. Informatore Fitopatol 50:43–46 Hohn T (2011) Caulimovirus. Caulimoviridae. In: The Springer index of Viruses. Springer, New York, pp 271–277. https://doi.org/10.1007/978-0-387-95919-1_41 Milusheva S, Rankova Z (2006) Serological identification of Plum pox virus in some economic important weeds. Selskostopanska Nauka (Agric Sci) 39:38–41 Vicchi V, Bellardi MG (1988) The role of weeds in the epidemiology of gladiolus viruses and MLO. Acta Hortic 234:371–378 Virscek Marn M, Mavric I, Urbancic-Zemljic M, Skerlavaj V (2004) Detection of Plum pox virus Potyvirus in weeds. Acta Hortic 657:251–254

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536

Cirsium vulgare (Bull thistle)

Cirsium vulgare (Bull thistle) Family: Asteraceae

Weed host

Alfalfa mosaic virus

(AMV)

Taxonomic position Genus: Alfamovirus

Family: Bromoviridae

AMV infection in plants of Cirsium vulgare was reported from New Zealand (Fletcher 2001). The virus-infected bull thistle plants do not exhibit obvious symptoms. The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Cirsium vulgare was reported from Italy and New Zealand (Vicchi and Bellardi 1988; Fletcher 1989). The virus-infected bull thistle plants do not exhibit external symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of CMV, refer to Cucumis sativus.

References Fletcher JD (1989) Additional hosts of Alfalfa mosaic virus, Cucumber mosaic virus and Tobacco mosaic virus in New Zealand. N Z J Crop Hortic Sci 17:361–362 Fletcher D (2001) New hosts of Alfalfa mosaic virus, Cucumber mosaic virus, Potato virus Y, Soybean dwarf virus, and Tomato spotted wilt virus in New Zealand. N Z J Crop Hortic Sci 29:213–217 Vicchi V, Bellardi MG (1988) The role of weeds in the epidemiology of gladiolus viruses and MLO. Acta Hortic 234:371–378

Cissus spp. Family: Vitaceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(INSV)

Citrullus lanatus (Watermelon)

537

INSV was reported to infect plants of Cissus spp. in Iran (Ghotbi and Shahraeen 2012; Ghotbi 2013). The virus-infected cissus plants exhibit symptoms of necrotic leaf spot, chlorosis, and stunting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sapinoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

C

References Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iran J Plant Pathol 49:41 Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses affecting propagated ornamentals in Northern Iran. Int Res J Microbiol 3:373–381

Citrullus colocynthis (Bitter apple) Family: Cucurbitaceae

Papaya ringspot virus Taxonomic position Genus: Potyvirus

Medicinal

(PRSV)

Family: Potyviridae

PRSV pathotype W infection in plants of Citrullus colocynthis was reported from Iran (Naeimifar et al. 2014). The virus-infected bitter apple plants exhibit mottling and chlorosis symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PRSV, refer to Carica papaya.

References Naeimifar M, Pourrahim R, Zadehdabagh G (2014) Natural infection of Citrullus colocynthis by Papaya ringspot virus-W in Iran. Plant Dis 98:1748

Citrullus lanatus (Watermelon) Synonyms Citrullus vulgaris Family: Cucurbitaceae

Edible fruit

Chilli leaf curl virus

(ChiLCV)

Taxonomic position Genus: Begomovirus

Family: Geminiviridae

538

Citrullus lanatus (Watermelon)

ChiLCV infection in plants of Citrullus lanatus was reported from Oman (Shahid et al. 2017). The virus-infected watermelon plants exhibit leaf curl as well as yellowing and crumpling of leaves. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of ChiLCV, refer to Capsicum annuum.

Chickpea chlorotic dwarf virus Taxonomic position Genus: Mastrevirus

(CpCDV)

Family: Geminiviridae

CpCDV infection in plants of Citrullus lanatus was reported from Tunisia and Morocco (Zaagueri et al. 2017; Radouane et al. 2019). The virus-infected watermelon fruits exhibit chlorotic mottling on rind and white hard portions inside the flesh. The virus is transmitted by leafhopper vectors in a persistent, circulative, and non-propagative manner. The virus is not transmissible by mechanical inoculation. For more details of CpCDV, refer to Cicer arietinum.

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

CGMMV infection in plants of Citrullus lanatus was reported from China, Japan, Korea, Australia, India, and also several Mediterranean countries (Komuro et al. 1968, 1971; Nagai et al. 1974; Lovisolo 1981; Lee et al. 1990; Varveri et al. 2002; Boubourakas et al. 2004; Shim et al. 2005; Yoon et al. 2008; Zhang et al. 2009; Zhao et al. 2013; Amer 2015; Shargil et al. 2017; Tesoriero et al. 2016; Sun et al. 2017). The typical symptoms of CGMMV include a mosaic patterning, chlorotic mottling, necrosis of pedicels, and fruit distortions (Komuro 1971), often resulting in a reduced yield and a lower market value (Shim et al. 2005). The virus is not transmitted by any insect vector (Vani and Varma 1993). The virus is spread by direct mechanical contact with a range of contaminated sources including soil and propagation stock materials. The virus is transmissible through seed and pollen (Wu et al. 2011). For more details of CGMMV, refer to Cucumis sativus.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Citrullus lanatus was reported from Serbia and Israel (Milojevic et al. 2012; Reingold et al. 2013; Xanthis et al. 2015). Early in the growing season, the virus-infected watermelon plants show severe symptoms including stunting, mosaic, mottling, blistering, and leaf curling with reduced leaf size, while those infected at later stages exhibited only a mild mosaic. The virus is transmitted by a number of aphid vectors in a non-persistent manner, and also by mechanical sapinoculation to number of host plants. For more details of CMV, refer to Cucumis sativus.

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Cucumber vein yellowing virus Taxonomic position Genus: Ipomovirus

(CVYV)

Family: Potyviridae

CVYV has been reported in plants of Citrullus lanatus from Almeria, Spain, Iran, Israel, Jordan, Turkey, Sudan, Syria, and Saudi Arabia (Louro et al. 2004; Bananej et al. 2006). The virus-infected watermelon plants exhibit symptoms of vein-clearing, and mild leaf chlorosis, with occasional splitting of fruits. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CVYV, refer to Cucumis sativus.

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(CABYV)

Family: Luteoviridae

CABYV infection in plants of Citrullus lanatus was reported from China, Algeria, Greece, Turkey, Lebanon, Spain, Italy, and Tunisia (Lecoq et al. 1994; Juarez et al. 2004; Hattab et al. 2005, 2009; Yardimci and Ozgonen 2007; Xiang et al. 2008). The virus-infected watermelon plants exhibit yellowing of the older leaves and leaves are thickened and brittle. Awide range of symptom intensity exists depending on the cultivars, varying from a yellowing limited to a few older leaves to a complete discoloration of plants (Lecoq et al. 1992). The virus infection does not affect fruit quality but rather induces flower abortions and reduces the number of fruit per plant. The virus is transmitted by aphid vectors, Myzus persicae, M. euphorbiae, and Aphis gossypii, in a circulative, non-propagative manner (Lecoq et al. 1992). The virus is not transmissible by mechanical sap-inoculation. For more details of CABYV, refer to Cucumis melo.

Cucurbit chlorotic yellows virus

(CCYV)

Taxonomic position CCYV is a tentative member of the genus Crinivirus and family Closteroviridae CCYV infection in plants of Citrullus lanatus was reported from China, Taiwan, Greece, and Japan (Huang et al. 2010; Okuda et al. 2010; Gu et al. 2011; Orfanidou et al. 2014). The virus-infected watermelon plants exhibit symptoms of chlorosis and interveinal chlorotic spots on lower leaves. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a semi-persistent manner. For more details of CCYV, refer to Cucumis melo.

Cucurbit leaf crumple virus

(CuLCrV)

Synonyms Cucurbit leaf curl virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

C

540

Citrullus lanatus (Watermelon)

CuLCrV infection in plants of Citrullus lanatus was reported from Florida and California, USA (Gilbertson 2002; Akad et al. 2008; Hagen et al. 2008; Turechek et al. 2010). The virus-infected watermelon plants exhibit yellowing, crumpling, and curling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not mechanically sap-transmissible. For more details of CuLCrV, refer to Cucumis melo.

Cucurbit yellow stunting disorder virus Taxonomic position Genus: Crinivirus

(CYSDV)

Family: Closteroviridae

CYSDV infection in plants of Citrullus lanatus was reported from Arizona, California and Florida (USA), Mexico, Portugal and UAE (Hassan and Duffus 1991; Kao et al. 2000; Louro et al. 2000; Brown et al. 2007; Kou et al. 2007; Polston et al. 2008; Turechek et al. 2010). The virus-infected watermelon plants exhibit stunting, deformation, interveinal chlorosis, and leaf mottling symptoms. The virus is transmitted by both the B and Q biotypes of B. tabaci in a semi-persistent manner (Berdiales et al. 1999; McGrath 2004). The virus is not mechanically sap-transmissible. For more details of CYSDV, refer to Cucumis melo.

Lettuce infectious yellows virus Taxonomic position Genus: Crinivirus

(LIYV)

Family: Closteroviridae

LIYV infection in plants of Citrullus lanatus was reported from Texas (USA) (Halliwell and Johnson 1992). The virus-infected watermelon plants exhibit various degrees of mosaic symptoms and fruits do not ripen completely and plants are stunted. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a semi persistent manner. The virus is not mechanically sap-transmissible. For more details of LIYV, refer to Lactuca sativa.

Melon chlorotic mosaic virus Taxonomic position Genus: Begomovirus

(MClMV)

Family: Geminiviridae

MClMV infection in plants of Citrullus lanatus was reported from Venezuela (Romay et al. 2015). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. A putative alphasatellite, MeCMVa1, was found associated with MeCMV in watermelon (Romay et al. 2010). For more details of MClMV, refer to Cucumis melo.

Melon necrotic spot virus Taxonomic position Genus: Gammacarmovirus

(MNSV)

Family: Tombusviridae

Citrullus lanatus (Watermelon)

541

MNSV infection in plants of Citrullus lanatus was reported from Japan, China, Crete, and Spain (Avgelis 1989, 1990; Ohki et al. 2008; Sela et al. 2013; Ruiz et al. 2016; Yu et al. 2016). The virusinfected watermelon plants in the field displayed necrotic symptoms on veins and leaves and sometimes on the petiole and cirrus. The fruits had yellowish brown necrosis, and the pericarp and flesh were watery and hollow. The virus is transmitted by the fungal vector Olpidium bornovanus and also by mechanical sap-inoculation to fewer than 3 plant families. The virus is transmissible by contact between plants. For more details of MNSV, refer to Cucumis melo.

Melon rugose mosaic virus Taxonomic position Genus: Tymovirus

(MRMV)

Family: Tymoviridae

Geographical distribution MRMV infection in plants of Citrullus lanatus was first reported from the People’s Democratic Republic of Yemen by Jones (1981). The virus spreads in Sudan and Yemen (Jones et al. 1986; Alhubaishi et al. 1987; Mahgoub et al. 1997). Symptoms and host(s) The virus-infected watermelon plants exhibit yellowing and rugose mosaic symptoms. The natural host range of this virus is limited to cucurbits. Transmission The virus is transmitted by beetle vectors in a semi-persistent manner. The virus is mechanically transmissible and systemic symptoms were observed only in some cucurbits, including watermelon, snake cucumber, melon, and cucumber. The virus was also found to be transmissible by leaf contact. The virus is transmissible through seed of melon and snake cucumber, but not seed of watermelon. Virion properties and genome The virions are non-enveloped, isometric capsid with T = 3 icosahedral symmetry, about 32 nm in diameter. The genome consists of a single molecule of positive-sense, ssRNA of MW c.2.1  106. The 30 -terminus has a tRNA-like structure (Jones et al. 1986; Martelli et al. 2002).

Melon severe mosaic orthotospovirus Taxonomic position Genus: Orthotospovirus

(MSMV)

Family: Tospoviridae

MSMV was detected in plants of Citrullus lanatus from Mexico (Ciuffo et al. 2009). The virus-infected watermelon plants exhibit mosaic and leaf deformation symptoms. The virus is transmitted by a thrips vector Frankliniella occidentalis in a persistent-propagative manner, and also by mechanical sapinoculation. For more details of MSMV, refer to Cucumis melo.

C

542

Citrullus lanatus (Watermelon)

Melon yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(MYSV)

Family: Tospoviridae

MYSV infection in plants of Citrullus lanatus was reported from Ecuador, Japan, and Taiwan (Chen et al. 2008; Okuda et al. 2002; Peng et al. 2011; Quito-Avila et al. 2014). The virus-infected watermelon plants exhibit chlorotic spotting symptoms. The virus is transmitted by thrips vectors in a persistent, propagative manner. For more details of MYSV, refer to Cucumis melo.

Moroccan watermelon mosaic virus Taxonomic position Genus: Potyvirus

(MWMV)

Family: Potyviridae

Geographical distribution MWMV infection was first reported in plants of Citrullus lanatus from Morocco by Fischer and Lockhart (1974). The virus spreads in Morocco, Africa, and the Mediterranean basin (Van der Meer and Garnett 1987; McKern et al. 1993; Lecoq et al. 2001). Symptoms and host(s) The virus-infected watermelon plants exhibit symptoms of mosaic, severe leaf chlorosis, and fruit deformations and necrosis. Transmission The virus is transmitted by the aphid vectors, Myzus persicae, Aphis gossypii, and Aphis spiraecola in the non-persistent manner (Owolabi and Ekpiken 2014; Chatzivassiliou et al. 2016). The virus is also mechanically sap-transmissible only to species belonging to the family Cucurbitaceae. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome consists of a molecule of linear, positive-sense single-stranded RNA of 9730 nt (EF579955 = NC_009995) (Baum et al. 1979; Van der Meer and Garnett 1987; Yakoubi et al. 2008; Revers and Garcia 2015; Wylie et al. 2017).

Papaya ringspot virus Taxonomic position Genus: Potyvirus

(PRSV)

Family: Potyviridae

PRSV infection in plants of Citrullus lanatus occurs worldwide (Halfeld-Vieira et al. 2004; Abdalla and Akhtar 2012; Ali et al. 2012; Lima et al. 2012; Romay et al. 2014; Nagendran et al. 2017). The virusinfected watermelon plants show, ringspots, mosaic, crinkling, and distortion symptoms on the leaves; and uneven skin surface of the fruits. The virus is spread by a number of different aphid species including the green peach and melon aphids in a non-persistent manner (Pearson and Liayanage 1997;

Citrullus lanatus (Watermelon)

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Turechek et al. 2010). The virus is also mechanically sap-transmissible. For more details of PRSV, refer to Carica papaya.

Pepo aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(PABYV)

C Family: Luteoviridae

PABYV infection in plants of Citrullus lanatus was reported from Greece (Lotos et al. 2016). The virusinfected watermelon plants exhibit severe interveinal yellowing and necrosis symptoms. The virus is transmitted by a number of aphid species in a circulative and non-propagative manner. The virus is not sap or seed-transmitted. For more details of PABYV, refer to Cucurbita pepo.

Squash leaf curl virus Taxonomic position Genus: Begomovirus

(SLCuV)

Family: Geminiviridae

SLCuV infection in plants of Citrullus lanatus was reported from Texas, USA, and Lebanon (Isakeit et al. 1994; Sobh et al. 2012). The virus-infected watermelon plants exhibit a bright yellow mosaic or mottle accompanied by leaf curling and severe stunting. The whitefly, Bemisia tabaci, transmits this virus in a circulative non-propagative manner. For more details of SLCuV, refer to Cucurbita pepo.

Squash mosaic virus Taxonomic position Genus: Comovirus

(SqMV)

Family: Secoviridae

SqMV infection in plants of Citrullus lanatus was reported from Western Samoa and Trinidad (Nelson et al. 1965; Pearson and Liayanage 1997; Chinnaraja et al. 2016). The virus-infected watermelon plants exhibit symptoms of severe stunting, systemic leaf mosaic, and mild leaf distortion. The virus is transmitted by beetle vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is seed-transmitted in Citrullus lanatus up to 13% (Lestari and Nurhayati 2014). For more details of SqMV, refer to Cucurbita pepo.

Squash vein yellowing virus Taxonomic position Genus: Ipomovirus

(SqVYV)

Family: Potyviridae

SqVYV was reported in plants of Citrullus lanatus from Puerto Rico, Florida, and also from Indiana (Egel and Adkins 2007; Adkins et al. 2007; Adkins et al. 2008; Baker et al. 2008; Turechek et al. 2010; Webb et al. 2012; Acevedo et al. 2013).

544

Citrullus lanatus (Watermelon)

The symptoms of SqVYV on watermelon start with a slight yellowing of the foliage followed by browning and the collapse of the entire vine within weeks of the first symptoms. These symptoms appear as the fruit is approaching harvestable size. The infected watermelon plants, often close to harvest, collapse as leaves and stems develop necrotic areas that spread, leading to death of the plant and fruit rind necrosis and discoloration, commonly known as watermelon vine decline. When cut open, the fruit often exhibit discolored blotches in the rinds. The flesh is often discolored (too red) and has an offtaste and flavor which make the fruit unmarketable. The whitefly vector, Bemisia tabaci, transmits the disease in a non-persistent manner; a minimum of 3 h is needed to complete a cycle of transmission by the whitefly vector (Webb et al. 2012). The virus is also mechanically sap-transmissible. For more details of SqVYV, refer to Cucurbita pepo.

Tobacco curly shoot virus Taxonomic position Genus: Begomovirus

(TbCSV)

Family: Geminiviridae

TbCSV infection in plants of Citrullus lanatus was reported from China (Zhao et al. 2017). The virusinfected watermelon plants exhibit severe leaf curling and yellowing symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of TbCSV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Citrullus lanatus was reported from North America, Australia, India, Iran, Japan, the Netherlands, Nigeria, and Yugoslavia (Pound 1949; Abdalla et al. 2012). The virus-infected watermelon plants exhibit ringspots and mosaic symptoms on the leaves and infected fruits develop prominent “pimples,” bumps and ringspots. The virus is transmitted by nematode vectors in a nonpersistent manner, and also through mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato leaf curl Palampur virus Taxonomic position Genus: Begomovirus

(ToLCPalV)

Family: Geminiviridae

ToLCPalV infection in plants of Citrullus lanatus was reported from Iran (Heydarnejad et al. 2013). The virus-infected watermelon plants exhibit leaf curl symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ToLCPalV, refer to Solanum lycopersicum.

Citrullus lanatus (Watermelon)

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Tomato leaf curl New Delhi virus

(ToLCNDV)

Synonyms Tomato leaf curl virus-India

Taxonomic position Genus: Begomovirus

C Family: Geminiviridae

ToLCNDV infection in plants of Citrullus lanatus was reported from Pakistan (Mansoor et al. 2000). The virus-infected watermelon plants exhibit symptoms of leaf curling, mottling, and stunted plant growth. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative nonpropagative manner. The virus is graft-transmissible. For more details of ToLCNDV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Citrullus lanatus was reported from North America, Japan, New Zealand, United Kingdom, Denmark, and Sweden (Provvidenti 1986). The virus-infected watermelon plants exhibit diffuse green yellow mosaic with some rugosity. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation, and by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Citrullus lanatus was reported from Tunisia (Mnari-Hattab et al. 2014). The virus-infected watermelon plants exhibit mild leaf curling with stunting; the fruits from infected plants become harden and when ripe show white pulp symptoms (white heart). The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. The virus is transmitted by grafting but not transmitted by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

Watermelon bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(WBNV)

546

Citrullus lanatus (Watermelon)

Geographical distribution WBNV was reported on Citrullus lanatus plants from Taiwan, Japan, and India (Singh and Krishnareddy 1996; Jain et al. 1998; Krupashankar 1998; Kunkalikar et al. 2011; Basavaraj et al. 2017; Holkar et al. 2017; Shruthi et al. 2017). Symptoms and host(s) The field symptoms of WBNV in watermelon initially develop as chlorotic mottling, yellow spots or patches, and mild crinkling of leaves. Subsequently, necrosis of buds in the growing tips results in dieback of veins. In the young crop, rapid dieback and wilting of plants develops dramatically causing a total loss in the affected plants. In the mature crop, shortened internodes, upright growth of younger shoots, necrosis on stem, petiole, and fruit stalk are commonly seen. Infected plants produce unmarketable small, deformed fruits with uneven surface, and necrotic or chlorotic rings, depending on the cultivar (Jain et al. 1998). Natural infection of WBNV was detected in chilli and tomato in North India (Kunkalikar et al. 2011; Holkar et al. 2017). Transmission The virus is successfully transmitted by the thrips vectors, Thrips palmi and T. flavus, in a persistentpropagative manner (Singh and Krishnareddy 1995; Shruthi et al. 2017). The virus multiplies in the vector and is retained when the vector molts. The virus is mechanically sap-transmissible and infects 22 host plant species belonging to 15 genera in six families comprising economically important crops and several weeds by mechanical sap-inoculations. Some hosts produced chlorotic local lesions, whereas others produced necrotic local lesions. The virus is not transmissible by contact between plants and is not transmitted by seed and pollen. Virion properties and genome The virions are spherical and membrane bound 80–100 nm in diameter. The genome comprised of three unique molecules of negative or ambisense ssRNA, designated L (large) 8916 nt (GU735408 = NC_038289), M (medium) 4794 nt (GU584185 = NC_038290), and S (small) 3401 nt (GU584184 = NC_038288) (Krupashankar 1998; Jain et al. 1998; Rajasekharam 2010; Li et al. 2011).

Watermelon chlorotic stunt virus Taxonomic position Genus: Begomovirus

(WmCSV)

Family: Geminiviridae

Geographical distribution WmCSV infection in plants of Citrullus lanatus was reported from Saudi Arabia, Sudan, Iran, Israel, Lebanon, Yemen, and Jordan (Bedford et al. 1994; Bananej et al. 1996, 1998, 2002; Kheyr-Pour et al. 2000; Antignus et al. 2003; Abudy et al. 2010; Al-Musa et al. 2011; Sufrin-Ringwald and Lapidot 2011; Khan et al. 2012; Ali-Shtayeh et al. 2012, 2014; Samsatly et al. 2012; Al-Saleh et al. 2014; DominguezDuran et al. 2018). Symptoms and host(s) The virus-infected watermelon plants exhibit vein yellowing, chlorotic mottling, curling of leaves, severe plant stunting, and a drastic reduction of fruit yield.

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The natural hosts of this virus include Citrullus lanatus, Cucumis melo, Cucumis melo var. flexuosus, Cucurbita moschata, Cucurbita pepo, wild cucurbits, Citrullus colocynthis, and Cucumis melo var. agrestis. Transmission WmCSV is transmitted by the whitefly Bemisia tabaci, the vector of begomoviruses in a circulative, non-propagative manner (Bananej et al. 2002; Ghanim et al. 2007; Kollenberg et al. 2014). Experimentally the virus has been transmitted to Datura stramonium, Phaseolus vulgaris and Nicotiana glutinosa by Agrobacterium-mediated inoculation of virus clones inducing leaf curling and a mild yellow mosaic (Bananej et al. 2002). The virus is not transmissible by mechanical inoculation but is transmissible by grafting. Virion properties and genome The structure of the virions of WmCSV has not been investigated. In common with all geminiviruses, the virions of WmCSV are likely geminate (twinned icosahedra). The genome of WmCSV is bipartite consisting of two circular, single-stranded DNA components. DNA-A contains 2745 nt (AJ012081 = NC_003708; AJ245650) and DNA-B of 2753 nt (AJ012082 = NC_003709; AJ245651) (Briddon 2001; Khan et al. 2012; Ali-Shtayeh et al. 2014; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the Old World the DNA A component of WmCSV encodes six genes, two in the virion-sense and four in the complementarysense, whereas the DNA B encodes one gene in each orientation. The expression and function of these genes has not been investigated for WmCSV (Dominguez-Duran et al. 2018).

Watermelon leaf mottle virus Taxonomic position Genus: Potyvirus

(WLMV)

Family: Potyviridae

Geographical distribution WLMV infection in plants of Citrullus lanatus was reported from the USA (Purcifull et al. 1998; De Sa et al. 2000). Symptoms and host(s) The virus-infected watermelon plants exhibit mottling and mosaic symptoms. Transmission The virus is transmitted by the aphid vector, Myzus persicae, in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome is a single molecule of linear, positive-sense single-stranded RNA. The complete genome sequence is not available; however, a partial genome sequence of 2316 nt (AF028004) at the 30 terminus is available (De Sa et al. 2000; Wylie et al. 2017).

C

548

Citrullus lanatus (Watermelon)

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

Geographical distribution WMV infection in plants of Citrullus lanatus was distributed worldwide including Central Valley of California (USA), Israel, Yugoslavia, Egypt, Spain, Italy, Tunisia, France, Brazil, Spain, Turkey, India, and Mediterranean countries (Bhargava and Joshi 1960; Bhargava et al. 1975; Lecoq 1992; Lecoq et al. 1998; Yuki et al. 2000; CABI/EPPO 2003; Halfeld-Vieira et al. 2004; Moradi 2011; Ali et al. 2012; Finetti-Sialer et al. 2012; Trkulja et al. 2014b; Kamberoglu et al. 2015). WMV isolates were previously classified into two distinct groups, namely, WMV-1 and WMV-2 (Ebrahimi-Nesbat 1974; Greber 1978; Purcifull and Hiebert 1979; Yeh et al. 1984). Those isolates failed to infect noncucurbitaceous plants, but infected cucurbitaceous plants were designated WMV-2, while the isolates infecting plants outside the cucurbitaceae were designated WMV-1, although the latter is now considered to be a strain of PRSV (PRSV-W). Symptoms and host(s) The virus-infected watermelon plants exhibit symptoms which differ according to the isolate and the host cultivar. On leaves, symptoms are mosaic, vein-banding and deformations of varying severity including blisters, filiformism, and important leaf size reduction. On fruits, severe discoloration and slight deformation are observed in some cultivar and with some isolates (Crescenzi et al. 2001). The virus has a broad host range in important crops, mostly among cucurbits but also pea, carrot, orchids (vanilla, Habenaria radiata). WMV also infects weed hosts. Transmission The virus is transmitted by at least 35 aphid species in 19 genera. Aphis craccivora, Aphis gossypii, and Myzus persicae are efficient vectors and transmit WMV in a non-persistent manner (Lecoq and Desbiez 2008). The virus is mechanically sap-transmissible and its host range was restricted mostly to Cucurbitaceae, Leguminosae, and Chenopodiaceae. The virus has not been reported to be seedtransmitted. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm in diameter. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 10,035 nt (AY437609 = NC_006262) (Baum 1980; Baum and Purcifull 1981; Desbiez and Lecoq 2004; Ali et al. 2006; Revers and Garcia 2015; Wylie et al. 2017).

Watermelon silver mottle orthotospovirus Synonyms Tomato spotted wilt virus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(WSMoV)

Citrullus lanatus (Watermelon)

549

Geographical distribution WSMoV infection in plants of Citrullus lanatus occurs worldwide (Iwaki et al. 1984; Kameya-Iwaki et al. 1988; Yeh et al. 1988, 1992; Honda et al. 1989; Pappu et al. 1998; Okuda et al. 2002; Rao et al. 2011, 2013; Yin et al. 2014, 2016). Symptoms and host(s) The virus-infected watermelon plants generally exhibit silver mottling on leaves and chlorotic mottling on fruits that are often malformed. Affected plants were severely stunted and had shortened internodes, upright growth of younger branches, tip necrosis, and dieback. Foliar symptoms included mottling, crinkling, yellow spotting, and narrowed leaf laminae. Fruits were small and malformed and exhibited necrotic spots or silver mottling. Fruit set was reduced. The virus has a very wide host range and infects the members of the Leguminosae, Solanaceae, Cucurbitaceae, and other families inducing necrotic and chlorotic local lesions, systemic wilting, necrosis, spotting, streaking, mosaic, mottling, leaf shape malformation, vein yellowing, ringspots, line patterns, yellow netting, and flower color-breaking are produced on various inoculated plants (Iwaki et al. 1984). Transmission The virus is transmitted by the thrips vectors, Thrips tabaci and T. palmi, in a persistent-propagative manner (Chen et al. 1990; Yeh et al. 1992). Virus is retained when the vector molts; multiplies in the vector (probably); and is not transmitted congenitally to the progeny of the vector. The virus is transmissible mechanically by sap-inoculation, and shows systemic symptoms in a range of economically important plants including melon, cucumber, sweet pepper, tomato, tobacco, and squash (Cucurbita pepo, C. maxima) (Iwaki et al. 1984). The virus is also transmissible by grafting but not transmissible by contact between plants and not transmissible by seed and pollen (EPPO/CABI 1996). Virion properties and genome The virions are spherical or pleomorphic, 80–120 nm in diameter. The genome comprises three unique molecules of negative or ambisense ssRNA, designated L (large) 8917 nt (AF133128 = NC_003832), M (medium) 4880 nt (U75379 = NC_003841), and S (small) 3534 nt (U78734 = NC_003843) (Yeh and Chang 1995; Yeh et al. 1996; Chu and Yeh 1998; Chu et al. 2001; Rao et al. 2013).

Watermelon virus A Taxonomic position Genus: Wamavirus

(WVA)

Family: Betaflexiviridae

Geographical distribution WVA infection in plants of Citrullus lanatus was reported from China (Xin et al. 2017). Symptoms and host(s) The virus-infected watermelon plant leaves show crinkling and mosaic symptoms. Virion properties and genome The genome is positive-sense single-stranded RNA of 8372 nt (KY363796 = NC_034377), excluding the poly (A) tail, and contains four open reading frames (ORFs). The largest ORF, ORF1 encodes a

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putative replication-associated polyprotein (RP) with three conserved domains. ORF2 and ORF4 encode a movement protein (MP) and coat protein (CP), respectively. The putative product encoded by ORF3, of an estimated molecular mass of 25 kDa, has no significant similarity with other proteins (Xin et al. 2017).

Zucchini yellow mosaic virus Taxonomic position Genus: Potyvirus

(ZYMV)

Family: Potyviridae

ZYMV infection in plants of Citrullus lanatus was reported from Turkey, southeastern California desert valleys (USA), Bosnia, and Herzegovina (Davis and Yilmaz 1984; Nameth et al. 1985; Pearson and Liayanage 1997; Halfeld-Vieira et al. 2004; Ozaslan et al. 2006; Ali et al. 2012; Vucurovic et al. 2012; Trkulja et al. 2014a). The virus-infected watermelon plants exhibit mottling, mosaic, and leaf filiformism symptoms. Fruits may present irregular coloration and slight to severe deformations. The virus is transmitted by aphid vectors such as Aphis gossypii and Myzus persicae in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ZYMV, refer to Cucurbita pepo.

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Plant Dis 101:1063 Shargil D, Smith E, Lachman O, Reingold V, Darzi E, Tam Y, Dombrovsky A (2017) New weed hosts for Cucumber green mottle mosaic virus in wild Mediterranean vegetation. Eur J Plant Pathol 148:473–480 Shim CK, Han KS, Lee JH, Bae DW, Kim DK, Kim HK (2005) Isolation and characterization of watermelon isolate of Cucumber green mottle mosaic virus (CGMMV-HY1) from watermelon plants with severe mottle mosaic symptoms. J Plant Pathol 21:167–171 Shruthi CR, Narabenchi GB, Devaraju G, Nirmala HR (2017) Extent of thrips infestation and Watermelon bud necrosis viral disease incidence in major watermelon growing areas of Karnataka. Int J Pure App Biosci 5(5):298–301 Singh SJ, Krishnareddy M (1995) Thrips flavus Scrank (Thysanoptera: Thripidae), a new insect vector of a Tospovirus infecting watermelon in India. Pest Manage Horticultural Ecosyst 1:115–118 Singh SJ, Krishnareddy M (1996) Watermelon bud necrosis: a new Tospovirus disease. Acta Hortic 431:68–77 Sobh H, Samsatly J, Jawhari M, Najjar C, Haidar A, Abou-Jawdah Y (2012) First report of Squash leaf curl virus in cucurbits in Lebanon. Plant Dis 96(8):1231 Sufrin-Ringwald T, Lapidot M (2011) Characterization of a synergistic interaction between two cucurbit-infecting begomoviruses: Squash leaf curl virus and Watermelon chlorotic stunt virus. Phytopathology 101:281–289 Sun Y, Niu X, Fan M (2017) Genome-wide identification of Cucumber green mottle mosaic virus-responsive microRNAs in watermelon. Arch Virol 162:2591–2602 Tesoriero LA, Chambers G, Srivastava M, Smith S, Conde B, Tran-Nguyen LTT (2016) First report of Cucumber green mottle mosaic virus in Australia. Aust Plant Dis Notes 11:1 Trkulja V, Josic Kovacic D, Mihic Salapura J, Stankovic I, Vucurovic A, Bulajic A, Krstic B (2014a) First report of Zucchini yellow mosaic virus in Watermelon in Bosnia and Herzegovina. Plant Dis 98:858 Trkulja V, Vasic J, Vukovic B, Stankovic I, Vucurovic A, Bulajic A, Krstic B (2014b) First report of Watermelon mosaic virus Infecting melon and watermelon in Bosnia and Herzegovina. Plant Dis 98:1749 Turechek WW, Kousik CS, Adkins S (2010) Distribution of four viruses in single and mixed infections within infected watermelon plants in Florida. Phytopathology 100:1194–1203 van der Meer FW, Garnett HM (1987) Purification and identification of a South African isolate of Watermelon mosaic virus – Morocco. J Phytopathol 120:255–270 Vani S, Varma A (1993) Properties of Cucumber green mottle mosaic virus isolated from water of river Jamuna. Indian Phytopath 46:118–122 Varveri C, Vassilakos N, Bem F (2002) Characterization and detection of Cucumber green mottle mosaic virus in Greece. Phytoparasitica 30:493–501 Vucurovic A, Bulajic A, Stankovic I, Ristic D, Nikolic D (2012) First report of Zucchini yellow mosaic virus in Watermelon in Serbia. Plant Dis 96:149 Webb SE, Adkins S, Reitz SR (2012) Semi-persistent whitefly transmission of Squash vein yellowing virus causal agent of viral watermelon vine decline. Plant Dis 96:839–844 Wu HJ, Qin BX, Chen HY, Peng B, Cai JH, Gu QS (2011) The rate of seed contamination and transmission of Cucumber green mottle mosaic virus in watermelon and melon. Sci Agric Sin 44:1527–1532

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Citrus spp. (Citrus aurantifolia; C. aurantium)

Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J GenVirol 98:352–354 Xanthis CK, Maliogka VI, Lecoq H, Dezbiez C, Tsvetkov I, Katis NI (2015) First report of Cucumber mosaic virus infecting watermelon in Greece and Bulgaria. J Plant Pathol 97(2):399 Xiang HY, Shang QX, Han CG, Li DW, Yu JL (2008) First report on the occurrence of Cucurbit aphid-borne yellows virus on nine cucurbitaceous species in China. Plant Pathol 57:390 Xin M, Zhang P, Liu W, Ren Y, Cao M, Wang X (2017) The complete nucleotide sequence and genome organization of a novel Betaflexivirus infecting Citrullus lanatus. Arch Virol 162(10):3239–3242 Yakoubi S, Desbiez C, Fakhfakh H, Wipf-Scheibel C, Marrakchi M, Lecoq H (2008) Biological characterization and complete nucleotide sequence of a Tunisian isolate of Moroccan watermelon mosaic virus. Arch Virol 153:117–125 Yardimci N, Ozgonen H (2007) First report of Cucurbit aphid-borne yellows virus in Turkey. Aust Plant Dis Notes 2:59 Yeh SD, Chang TF (1995) Nucleotide sequence of the N Gene of Watermelon silver mottle virus, a proposed new member of the genus Tospovirus. Phytopathology 85:58–64 Yeh SD, Gonsalves D, Provvidenti R (1984) Comparative studies on host range and serology of Papaya ringspot virus and Watermelon mosaic virus 1. Phytopathology 74(9):1081–1085 Yeh SD, Cheng YH, Jih CL, Chen CC, Chen MJ (1988) Identification of Tomato spotted wilt virus infecting horn melon and watermelon. Plant Prot Bull 30:319–320 Yeh SD, Lin YC, Cheng YH, Jih CL, Chen MJ, Chen CC (1992) Identification of Tomato spotted wilt-like virus on watermelon in Taiwan. Plant Dis 76:835–840 Yeh S-D, Sun I-J, Ho H-M, Chang T-F (1996) Molecular cloning and nucleotide sequence analysis of the SRNA of Watermelon silver mottle virus. Acta Hortic 431:244–260 Yin YY, Fang Q, Lu X (2014) Detection of Watermelon silver mottle virus infecting watermelon in Yunnan, southwest of China. J Plant Pathol 96:S4.123 Yin Y-Y, Lu X, Li T-T, Li H-X, Guo M, Zhao L-L, Zhang X-Y, Ding M (2016) Identification of Watermelon silver mottle virus infecting watermelon in Yunnan. Acta Phytopathol Sinica 46(4):461–468 Yoon JY, Choi GS, Choi SK, Hong JS, Choi JK, Kim W, Lee GP, Ryu KH (2008) Molecular and biological diversities of Cucumber green mottle mosaic virus from cucurbitaceous crops in Korea. J Phytopathol 156:408–412 Yu C, Wang D, Zhang X, Shi K, Li X, Yuan X (2016) First report of Melon necrotic spot virus in watermelon in China. Plant Dis 100:1511 Yuki VA, Rezende JAM, Kitajima EW, Barroso PAV, Kuniyuki H, Groppo GA, Pavan MA (2000) Occurrence, distribution and relative incidence of five viruses infecting cucurbits in the State of Sao Paulo, Brazil. Plant Dis 84:516–520 Zaagueri T, Mnari-Hattab M, Zammouri S, Hajlaoui MR, Accotto GP, Vaira AM (2017) First report of Chickpea chlorotic dwarf virus in watermelon (Citrullus lanatus) in Tunisia. Plant Dis 101:392 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zhang YJ, Li GF, Li MF (2009) Occurrence of Cucumber green mottle mosaic virus on cucurbitaceous plants in China. Plant Dis 93:200 Zhao H-R, Lin Z-Y, Zhu J-Z, Zhang Y-D, Gao B-D (2013) First report of Cucumber green mottle mosaic virus (CGMMV) in Hunan Province. Acta Phytopathol Sinica 43(2):219–221 Zhao LL, Ding M, Zhang XY, Yin YY, Li TT, Zhang ZK (2017) First report of Tobacco curly shoot virus (TBCSV) and its associated satellites from watermelon in China. J Plant Pathol 99:761–764

Citrus spp. (Citrus aurantifolia; C. aurantium; C. clementina; C. decumana; C. grandis; C. jambhiri; C. limon; C. medica; C. paradisi; C. reticulata; C. sinensis; C. unshia; C. volkameriana) Family: Rutaceae

Edible fruit

Apple stem grooving virus Synonyms Citrus tatter leaf virus

(ASGV)

Citrus spp. (Citrus aurantifolia; C. aurantium)

Taxonomic position Genus: Capillovirus

557

Family: Betaflexiviridae

ASGV infection in plants of Citrus spp. was first discovered in the United States in 1962 (Wallace and Drake 1962). The virus spreads in the Eastern Asian region, the Pacific region, Australia, China, Japan, Korea, South Africa, Taiwan, Italy and the USA (Zhang et al. 1988; Ke and Wu 1991; Broadbent et al. 1994; Kawai et al. 1996; Hailstones et al. 2000; Ito et al. 2002a, b, 2003; Lovisolo et al. 2003). The virus-infected citrus plants are often symptomless. Chlorotic leaf symptoms are produced in Citrus excelsa, Rusk and Troyer citranges, and other trifoliate orange hybrids. Leaves of C. excelsa may be deformed, but infected plants often recover after the initial reaction. Stems of citrange plants may be deformed and have a zigzag growth pattern associated with chlorotic areas on the stem. Citrange and citrange hybrids are often pitted. The virus-affected trifoliate orange plants are stunted and chlorotic. Over blooming and early maturation of fruit are observed for a few years and the trees may die within 5–6 years after grafting. No natural vector is known for this virus. The virus is transmissible by mechanical sap-inoculation to herbaceous hosts (Nishio et al. 1982). The virus is seed-transmitted in certain Citrus spp. (Tanner et al. 2010). The virus is also transmitted by grafting. The virus spreads by the use of infected budwood (Nishio et al. 1989; Ohira et al. 1995). For more details of ASGV, refer to Malus domestica.

Citrus bark cracking viroid

(CBCVd)

Synonyms Citrus viroid IV Taxonomic position Genus: Cocadviroid

Family: Pospiviroidae

Geographical distribution CBCVd infection in different Citrus spp. is distributed in the major production areas worldwide viz., Sudan, China, Israel, Iran, Turkey, Sicily, Brazil, Japan, China, Spain, the USA, Italy and South Africa (Hadas et al. 1989; Onelge et al. 2000; Ito et al. 2002a, b; Malfitano et al. 2002; Semancik and Vidalakis 2005; Kunta et al. 2007; Eiras et al. 2009; Mohamed et al. 2009; Cao et al. 2010; Cook et al. 2012; Al-Harthi et al. 2013; Murcia et al. 2015; Rizzo et al. 2017). Symptoms and host(s) The viroid-infected trifoliate orange (Citrus trifoliata, syn. Poncirus trifoliata) plants exhibits bark cracking symptoms. Citrus medica plants exhibit mild and transient symptoms, usually leaf epinasty. C. sinensis plants initially exhibit, leaf bend of a single leaf and later, petiole browning developed on all test plants (Murcia et al. 2015). Transmission The viroid is transmitted by grafting, cuttings, and mechanical means. The viroid is transmissible mechanically to Benincassa hispida (Puchta et al. 1991). Etiology and genome properties The genome consists of single-stranded circular RNA of 284 nt, which is non-coding (X14638 = NC_003539) (Puchta et al. 1991; Gora-Sochacka 2004; Giguere et al. 2014).

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Citrus bent leaf viroid

Citrus spp. (Citrus aurantifolia; C. aurantium)

(CBLVd)

Synonyms Citrus viroid I

Taxonomic position Genus: Apscaviroid

Family: Pospiviroidae

Geographical distribution CBLVd was first identified in1988 and termed as CbVd-Ia and Ib based on the distinct mobility in sequential-PAGE analysis (Duran-Vila et al. 1988). The viroid infection in Citrus spp. plants occur worldwide including Australia, Israel, Iran, the United States, Uruguay, Japan, the Philippines, Italy, China, Pakistan, Sudan, Malaysia, European countries and AUE (Ashulin et al. 1991; Semancik et al. 1997; Hataya et al. 1998; Ito et al. 2002a, b; Alavi et al. 2005; Wang et al. 2008a, b; Cao et al. 2009; Mohamed et al. 2009; Eiras et al. 2009; Bagherian and Izadpanah 2011; Pagliano et al. 2013; Al-Harthi et al. 2013; Al-Shariqi et al. 2013; Mazhar et al. 2014; Khoo et al. 2017; Rizzo et al. 2017).

Symptoms and host(s) The viroid-infected Etrog citron plants display leaf bending and point necrosis on the leaf midribs. A Valencia/trifoliate citrus planting inoculated with pure isolates of CBLVd (CVd-1a type exactly) show specific reactions of “Trifoliate deep pit” consisting of deep pits in the wood of the trifoliate trunk with corresponding pointed pegs in the bark which is associated with a significant reduction of tree canopy (Roistacher et al. 1993). The viroid infection causes a necrotic lesions on the underside of the leaf mid-vein and/or bent leaf symptoms, a descriptive term for the viroid, on Arizona 861-S-1 “Etrog citron”(Citrus medica) (Duran-Vila et al. 1993; Verniere et al. 2004). The host range is restricted almost exclusively to Rutaceous hosts.

Transmission The viroid is transmissible by grafting and also mechanically through contaminated implents. Use of infected budwood helps in the spread of the disease.

Etiology and genome properties The genome consists of single-stranded circular RNA. Considerable variability in the nucleotide sequence was found in CBLVd isolates (Ben-Shaul et al. 1995). CbVd-1a consists of 327-nt (Semancik et al. 1997) and CbVd-1b consists of 318-nt (Ashulin et al. 1991). The isolates in Japan and the Philippines consisted of 328 nt which is 10-nt larger than the type CBLVd-225A in Israel. The Philippines sample also contained a sequence variant of 329-nt. An isolate from China consisted of 318-nt and shared 98% identity to CBLVd-225A. CVd-Ia was separated from CVd-Ib by two 5-nt insertions located in upper (50 -AGCUG-30 ) and the lower (50 -CUUCU-30 ) strand of the right terminal region. Since both of the 5-nt insertions are similar to the adjacent sequences (50 -AGUUG-30 and 50 -CUUCU-30 ), it was hypothesized that CVd-Ia is a derivative of Vd-Ib caused by partial sequence duplications and substitutions taking place in the right terminal region (Hataya et al. 1998). The reference sequence (U21125 = NC_001651) has 315 nt (Gora-Sochacka 2004; Giguere et al. 2014).

Citrus spp. (Citrus aurantifolia; C. aurantium)

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Citrus chlorotic dwarf associated virus Taxonomic position Genus: Unassigned

(CCDaV)

Family: Geminiviridae

Geographical distribution CCDaV infection in plants of Eureka lemon (Citrus limon) was reported from China (Guo et al. 2015; Zhou et al. 2017b). Symptoms and host(s) The virus-infected citrus plants exhibit symptoms of mosaic, distortion, and shrinkage on tender shoots and young leaves. The branches of the infected trees show shortened internodes and narrowed leaf blades. In the first year of infection, the trees could flower and bear fruit normally, but later flowering and fruit bearing were significantly reduced, indicating loss of vitality of the trees. Transmission The virus is suspected to be transmitted by the citrus whitefly vector, Dialeurodes citri. The virus is transmissible by grafting. Virion properties and genome The virions are twin isometric morphology. The genome is single-stranded circular DNA consists of 3640 nts (JQ920490 = NC_018151) (Howd et al. 2002; Loconsole et al. 2012b; Guo et al. 2015; Zerbini et al. 2017).

Citrus chlorotic spot dichorhavirus Taxonomic position Genus: Dichorhavirus

(CiCSV)

Family: Rhabdoviridae

Geographical distribution CiCSV infection in plants of Citrus sinensis was reported from Brazil (Chabi-Jesus et al. 2018). Symptoms and host range The virus-infected citrus plants exhibit chlorotic spot symptoms. Transmission The virus is transmitted by a mite vector, Brevipalpus yothersi in a persistent-propagative manner. The virus is transmitted by mechanical sap-inoculation. Virion properties and genome The virions are bacilliform, non-enveloped and measures 40  100110 nm. The genome is composed of two single-stranded negative-sense RNAs. RNA1 consists of 6518 nt (KY700685) and RNA2 is 5987 nt (KY700686) (Walker et al. 2018).

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Citrus spp. (Citrus aurantifolia; C. aurantium)

Citrus coguvirus Synonyms Citrus concave gum-associated virus (CCGaV) Taxonomic position Genus: Coguvirus

Family: Unassigned

Geographical distribution The virus infection in plants of Citrus sinensis was reported from Italy (Navarro et al. 2018a, b). Virion properties and genome The virions are flexuous, nonenveloped particles. The genome consists of two segments of linearnegativesense single-stranded RNAs (with RNA2 being ambisense). RNA1: 6681 nt (KX960112); RNA2: 2703 nt (KX960111). RNA1 encodes the RdRp in the viral complementary (vc) strand, while RNA2 encodes a putative MP in the viral strand, and nucleoprotein (NP) in the vc strand (Navarro et al. 2018a, b).

Citrus dwarfing viroid

(CDVd)

Synonyms Citrus viroid III Taxonomic position Genus: Apscaviroid

Family: Pospiviroidae

Geographical distribution CDVd was first reported as CVd-IIIa and IIIb (Duran-Vila et al. 1988). CDVd infection in Citrus spp. plants occurs worldwide including Japan, Tunisia, California, Uruguay, Brazil, Spain, Italy, Sudan, New Zealand, China, Syria and Corsica (Ito et al. 2002a, b; Malfitano et al. 2002; Elleuch et al. 2006; Kunta et al. 2007; Rizza et al. 2007; Wang et al. 2008a, b; Cao et al. 2009; Eiras et al. 2009, 2010, 2013; Mohamed et al. 2009; Quemin et al. 2011; Al-Harthi et al. 2013; Pagliano et al. 2013; Murcia et al. 2015; Abou Kubaa et al. 2016; Najar et al. 2017; Rizzo et al. 2017). Symptoms and host(s) CVd-III has a narrow host range and causes specific symptoms in “Etrog citron”, an indicator species for citrus viroids, characterized by petiole ringing, necrosis, and leaf drooping as a result of petiole bending. The presence of CVd-IIIb significantly reduced both the trunk section of the scion and rootstock, and also the canopy volume. For example, the presence of CVd-IIIb alone caused a 47% reduction of scion cross-sectional area and a 50% reduction in tree volume in “Valencia” sweet orange trees grafted on trifoliate orange rootstock (Roistacher et al. 1993). Because of this interesting characteristic, selections of CVd-III variants were tested as dwarfing agents for citrus cultivation (Broadbent et al. 1986; Bar-Joseph 1993; Gillings et al. 1991; Semancik et al. 1997; Villalobos et al. 1997; Owens et al. 2000; Hutton et al. 2000). Because of this use, it was proposed to rename CVd-III as Citrus dwarfing viroid (CDVd) (Flores et al. 1998; Owens et al. 1999; Verniere et al. 2004; Murcia et al. 2015).

Citrus spp. (Citrus aurantifolia; C. aurantium)

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Transmission The viroid is mechanically sap-transmissible and also through contaminated tools. Use of infected budwood and root stocks is the primary mode of spread. Etiology and genome properties The genome consists of single-stranded circular RNA of 292~297 nt, and sequence changes were confined largely to the lower portions of the central conserved region and variable domain. The variants of CVd-IIIa type (297 nt) and CVd-IIIb type (294 nt) were predominant in the field isolates collected from Israel and Costa Rica (Owens et al. 2000). The Tunisian isolates of CVd-III consist of 293–297 nt and showed nucleotide changes in 26 positions out of 297 (8.8%) compared to an Australian isolate (Elleuch et al. 2006). The reference sequence (AF447788 = NC_003264) has 292 nt (Gora-Sochacka 2004; Giguere et al. 2014).

Citrus exocortis viroid Taxonomic position Genus: Pospiviroid

(CEVd)

Family: Pospiviroidae

Geographical distribution CEVd is present in most Citrus spp. growing areas where susceptible rootstock is used. It is widespread in Italy, South America (Brazil, Uruguay and Argentina), Asia (China, India, Pakistan, Japan), Australia, Azerbaijan, Syria and the Mediterranean region (Spain), Tunisia, but is of limited occurrence in the USA and N. Africa (Nariani et al. 1968; Semancik and Weathers 1972; Semancik 1988; Semancik et al. 1993; Ito et al. 2002a, b; Malfitano et al. 2002; Ahlawat and Pant 2003; Ramachandran et al. 2003; Arif et al. 2005; Ragozzino et al. 2005; Elleuch et al. 2006; Kunta et al. 2007; Rizza et al. 2007; Wang et al. 2008a, b; Eiras et al. 2009, 2010, 2013; Mohamed et al. 2009; Abubaker and Elhassan 2010; Al-Harthi et al. 2013; Pagliano et al. 2013; Mazhar et al. 2014; Murcia et al. 2015; Abou Kubaa et al. 2016; Tan et al. 2016; Najar et al. 2017; Rizzo et al. 2017). Symptoms and host(s) The viroid-infected citrus remains symptomless in most scion cultivars. Symptoms are expressed only when infected scions are grafted on sensitive rootstocks, such as trifoliate orange, citranges and “Rangpur” lime. Between 1 and 2 years after graft inoculation, plants of Citrus sinensis (sweet orange) on Citrus trifoliata (=Poncirus trifoliata; trifoliate orange) show classical symptoms of bark shelling and scaling of the rootstock and a decrease in tree vigour leading to stunting and yield reduction. Citrus medica (Etrog citron), an indicator species, show leaf epinasty and rugosity cracking and browning of the underside of the veins and severe stunting visible from 3 weeks to 6 months after graft or razor slash inoculation (Bitters et al. 1987; Verniere et al. 2004; Murcia et al. 2015). The host range is largely restricted to the citrus family (Rutaceae) although some species in the Solanaceae (Solanum tuberosum, S. lycopersicum, Petunia hybrida) and Compositae (Gynura aurantiaca, G. saramentosa) are susceptible. Transmission There is no evidence for transmission by a vector. Tests for seed transmission in citrus have been negative except for a single report which might be reinterpreted as natural spread. The viroid is transmissible easily by mechanical sap-inoculation of sap from Gynura but with difficulty from Citrus (Garnsey and Whidden 1973). Transmission from citrus to citrus and from citrus to petunia was

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Citrus spp. (Citrus aurantifolia; C. aurantium)

accomplished by means of Cuscuta subinclusa. The viroid is easily spread on infected budwood and contaminated propagation tools (Garnsey 1967). Etiology and genome properties The genome consists of single-stranded circular RNA of 371 nt (J02053 = NC_001464) (Gross et al. 1982; Visvader et al. 1982; Onelge 1997; Rakowski et al. 1994; Elleuch et al. 2003; Gora-Sochacka 2004; Murcia et al. 2011; Giguere et al. 2014).

Citrus leaf blotch virus

(CLBV)

Synonyms Dweet mottle virus Taxonomic position Genus: Citrivirus

Family: Betaflexiviridae

Geographical distribution CLBV has been detected in several Citrus spp. from Australia, the USA (Florida and California), Italy, France, Japan, New Zealand, Cuba, China and Spain (Galipienso et al. 2001, 2004; Vives et al. 2002, 2008a; Krueger et al. 2005; Guardo et al. 2007; Harper et al. 2008; Hernandez-Rodriguez et al. 2016; Cao et al. 2017b). Symptoms and host(s) The virus-infected citrus plants exhibit chlorotic leaf blotching in dweet tangor and stem pitting in Etrog Citron and bud union crease when propagated on Carrizo citrange (Galipienso et al. 2001; Vives et al. 2002). The virus has not been found naturally infecting hosts other than citrus. Transmission The virus is sap-transmissible when partially purified virus was used for inoculation. The virus is seedtransmitted in Carrizo citrange, Nagami kumquat or sour orange (Citrus aurantium) at a rate of 2.5% (Guerri et al. 2004). The primary spread of the virus takes place by the use of infected bud sticks. Vector transmission is not demonstrated. Virion properties and genome The virions are filamentous particles of about 960  14 nm in size, contains a single positive-sense RNA, encapsidated by a 42 kDA coat protein (Galipienso et al. 2001). The gnomic RNA has 8747 nt (AJ318061 = NC_003877) organized in three open reading frames (ORFs) and untranslated regions (UTR) of 73 and 541 nt at the 50 and 30 termini, respectively (AJ318061) (Vives et al. 2001, 2008b; Adams et al. 2004; Hajeri et al. 2010).

Citrus leaf rugose virus Taxonomic position Genus: Ilarvirus

(CiLRV)

Family: Bromoviridae

Citrus spp. (Citrus aurantifolia; C. aurantium)

563

Geographical distribution CiLRV infection in plants of Citrus spp. was reported from Argentina, Japan and Florida (USA) (Fulton 1981; Frison and Taher 1991). Symptoms and host(s) The virus infects a wide range of citrus hosts. The virus-infected Mexican lime plants develop leaf puckering, Eureka lemon develops pinpoint chlorotic spotting and Duncan grapefuit shows stunting and chlorosis. Transmission The virus is mechanically sap-transmissible to a number of citrus species and also to herbaceous plants which are local lesion hosts. The virus is transmissible through contaminated cutting tools. The virus is graft-transmissible and no vector is known. No seed transmission is observed. Virion properties and genome The virions are isometric varying from 25 to 32 nm in diameter. The genome is tripartite single-stranded RNA (Garnsey 1975; Ge and Scott 1994; Scott and Ge 1995a, b). RNA1= 3404 nt (NC_003548); RNA2= 2990 nt (NC_003547); and RNA3= 2289 nt (NC_003546). A subgenomic RNA4 (1009 nt) that codes for the coat protein is also present (Scott 2011a, b).

Citrus leprosis virus C Taxonomic position Genus: Cilevirus

(CiLV-C)

Family: Kitaviridae

Geographical distribution CiLV-C infection in plants of Citrus spp. was reported from Brazil, Argentina, Paraguay, Uruguay, Honduras, Venezuela, Costa Rica, Panama, Honduras, Guatemala, Bolivia, Mexico, Colombia, Belize and the USA (Dominguez et al. 2001; Saavedra et al. 2001; Childers et al. 2003; Gomez et al. 2005; Leon et al. 2006; Rodrigues et al. 2007; CABI/EPPO 2013; Castillo et al. 2011; Izquierdo-Castillo et al. 2011; Hartung et al. 2015). Symptoms and host(s) The virus-infected citrus plants exhibit round to elliptical local lesions on fruits, leaves and twigs. The severity of the lesions varies with the type of citrus and the region of origin. Leaf symptoms are usually round with a dark-brown central spot about 2–3 mm diameter, surrounded by a chlorotic halo, in which 1–3 brownish rings frequently appear surrounding the central spot; the overall lesion size varies from 10 to 30 mm, though larger lesions may form by the fusion of two or more adjacent lesions. On green fruits, the lesions are initially yellowish, becoming more brown or black, sometimes depressed, and reducing the market value of the fruits. On stems, lesions may be protuberant, cortical, grey or brown. Lesions may coalesce when present in large numbers, leading to girdling and death of the twig (Bastianel et al. 2006; Melzer et al. 2012). The virus is found primarily on sweet orange, but sour orange and mandarins are also susceptible, although other citrus cultivars do not normally show conspicuous symptoms.

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564

Citrus spp. (Citrus aurantifolia; C. aurantium)

Transmission This virus is transmitted by mite vectors, Brevipalpus californicus, B. obovatus, B. yothersi and B. phoenicis have been reported, but only B. phoenicis has proven to be an efficient vector (Rodrigues et al. 2000, 2003, 2005; Childers et al. 2001; Nunes et al. 2012; Roy et al. 2015c; GarciaEscamilla et al. 2018). The virus is also sap-transmissible from citrus to citrus and to herbaceous plants, mainly those in the family Chenopodiaceae. The virus is transmissible by grafting; not transmissible by seed and pollen (Colariccio et al. 1995; Childers and Rodrigues 2011). Virion properties and genome The virions are short, bacilliform, 100–110 nm long and 40–50 nm wide (Knorr 1968; Kitajima et al. 1972). The genome consists of two molecules of linear, positive-sense, ssRNA: RNA1 consists of 8745 nt (DQ352194 = NC_008169) and RNA2 of 4986 nt (DQ352195 = NC_008170). The RNA molecules are polyadenylated at the 30 -termini, and contain a “cap” structure in the 50 -termini (LocaliFabris et al. 2006; Pascon et al. 2006).

Citrus leprosis virus C2 Taxonomic position Genus: Cilevirus

(CiLV-C2)

Family: Kitaviridae

Geographical distribution CiLV-C2 infection in plants of Citrus spp. was reported from Colombia (Roy et al. 2013). Symptoms and host(s) The virus-infected plants exhibit round to elliptical local lesions on fruits, leaves and twigs. Leaf symptoms are usually round with a dark-brown central spot. On green fruits, the lesions are initially yellowish, becoming more brown or black, sometimes depressed, and reducing the market value of the fruits. On stems, lesions may be protuberant, cortical, grey or brown. Lesions may coalesce when present in large numbers, leading to girdling and death of the twig. Transmission The virus is transmitted by the false spider mite or flat mite, Brevipalpus phoenicis (Roy et al. 2015c). Virion properties and genome The virions are approximately 105  45 nm in diameter. The genome consists of two molecules of linear, positive-sense, ssRNA of about 8717 nt (RNA1; JX000024 = NC_038848) contained two open reading frames (ORFs) and 4989 nt (RNA2; JX000025 = NC_038849) contained five ORFs that encode the movement protein (MP) and four hypothetical proteins (p7, p15, p24, and p61) (Roy et al. 2013).

Citrus leprosis N dichorhavirus Synonyms Citrus necrotic spot virus

(CiLV-N)

Citrus spp. (Citrus aurantifolia; C. aurantium)

Taxonomic position Genus: Dichorhavirus

565

Family: Rhabdoviridae

Geographical distribution The nuclear type of Citrus leprosis virus (CilV-N) has been reported from Brazil, Panama, Colombia, and Mexico (Bastianel et al. 2010; Kitajima et al. 2011; Cruz-Jaramillo et al. 2014; Roy et al. 2014, 2015a, b; Hartung et al. 2015; Ramos-González et al. 2017). Symptoms and host(s) The virus affects citrus species without spreading systemically and causes localized lesions on leaves, stems, fruit, and twigs. Lesions initially appear chlorotic, typically developing a necrotic center. Lesions on fruit later develop into necrotic ringspots with a sunken center, or on twigs as necrotic lesions which may coalesce and girdle twigs. The lesions are centered on feeding sites of the tenuipalpid mite vectors. Transmission The virus spreads naturally by tenuipalpid mites of the genus Brevipalpus (Brevipalpus phoenicis; B. californicus sensu lato), and can also be transmissible by tip grafting of symptomatic shoots (Knorr 1968; Bastianel et al. 2010; Roy et al. 2015c; Garcia-Escamilla et al. 2018). Virion properties and genome The virions are bacilliform or bullet-shaped, of c.20–30 nm by 120–135 nm, with most particles accumulating in the nucleus. The genome is composed of two single-stranded negative-sense RNAs of 6307 nt (RNA1: KY751404) and 5831 nt (RNA2: KY751405). RNA1 encodes five ORFs, whereas RNA2 encodes the RdRp (Cruz-Jaramillo et al. 2014; Ramos-Gonzalez et al. 2017; Walker et al. 2018).

Citrus psorosis ophiovirus

(CPsV)

Synonyms Citrus ring spot virus Taxonomic position Genus: Ophiovirus

Family: Aspiviridae

Geographical distribution CPsV was first reported on Citrus spp. in Florida by Fawcett (1933). The virus spreads in Argentina, Australia, France, Greece, Iran, Italy, Spain, Turkey, Syria, Uruguay, North and South America, South Africa, Japan, New Zealand, India and the Mediterranean basin (Derrick et al. 1988; Ahlawat 1989; Roistacher 1993; Pant et al. 1997; Alioto et al. 2000; Loconsole et al. 2002; Tessitori et al. 2002; Loconsole et al. 2009; Ito et al. 2011; Quemin et al. 2011; Abou Kubaa et al. 2014; Rizzo et al. 2017). Psorosis A and B strains are reported on different Citrus spp. Symptoms and host(s) Citrus species infected with CPsV exhibit symptoms like epinasty, chlorotic flecks, leaf mottling, ringspots or large, irregular chlorotic patterns on mature leaves. Bark scaling of trunk and main branches and depending on strain (psorosis B) symptoms may include rampant bark scaling even on small limbs and twigs. The lesions enlarge progressively and will cover the trunk and main branches.

C

566

Citrus spp. (Citrus aurantifolia; C. aurantium)

Bark scaling is mainly a disease of sweet orange, mandarin and grapefruit, the most susceptible being sweet orange. The other species like sour orange (C. aurantium), lemon (C. limon), rough lemon (C. jambhiri) and Pummelo (C. grandis) will carry the disease without showing bark scaling (Roistacher 1991, 1993) but symptoms on the young leaves have been observed. Gum may accumulate below the scales and may impregnate the xylem producing wood staining and vessel occlusion (Roistacher 1993). Transmission Virus transmission by a vector has not been definitively demonstrated, but CPsV has been detected in zoospores of an Olpidium-like fungus from the roots of infected trees (Palle et al. 2005). An aerial vector has been suspected in Argentina (e.g. Pujol and Benatena 1965). The virus is transmissible by mechanical sap-inoculation to herbaceous test plants like Chenopodium quinoa, C. amaranticolor, and Gomphrena globosa, but sap-transmission is often erratic and not reliable for routine tests. The virus is transmitted by grafting. The virus is seed-transmitted ranging between 1–19% in the citranges (C. sinensis x P. trifoliata) and trifoliate orange (Childs and Johnson 1966; Campiglia et al. 1976; D’Onghia et al. 2000). In most citrus areas, psorosis seems to spread only through propagation of infected buds (Manjunath 1981). Virion properties and genome The virions are non-enveloped, naked filamentous nucleocapsids about 3 nm in diameter, forming kinked circles of at least two different contour lengths (300–500 and 1500–2500 nm). The circles (open form) can collapse to form pseudo-linear duplex structures of 9–10 nm in diameter (collapsed form) (Garcia et al. 1994; Navas-Castillo and Moreno 1995; Barthe et al. 1998). The genome is linear, negative-sense single-stranded RNA and consists of three segments. RNA1 (8186 nt AY654892 = NC_006314) is contained in the B component; and RNAs 2 (1645 nt AY654893 = NC_006315) and 3 (1447 nt AY654894= NC_006316) are packaged in the P component (Sanchez de la Torre et al. 1998, 2002; Naum-Ongania et al. 2003; Martin et al. 2005; Milne et al. 2011; Garcia 2012).

Citrus sudden death-associated virus Taxonomic position Genus: Marafivirus

(CSDaV)

Family: Tymoviridae

Geographical distribution CSDaV infection in plants of Citrus spp. was reported from Brazil (Roman et al. 2004; Maccheroni et al. 2005). Symptoms and host(s) The virus has no known host other than citrus. Transmission No vector is identified for this virus. The virus is not mechanically sap-transmissible. The virus is transmissible by grafting and is disseminated over long distances through the transport of infected propagating material (Roman et al. 2004).

Citrus spp. (Citrus aurantifolia; C. aurantium)

567

Virion properties and genome The virions are non-enveloped, isometric capsid with T = 3 icosahedral symmetry, about 30 nm in diameter. The genome is a monopartite, linear, positive-sense, single-stranded RNA of 6820 nt (AY884005 = NC_006950). There is no VPg at the 50 -terminus (Martelli et al. 2002; Bassanezi et al. 2003; Maccheroni et al. 2005; Hammond et al. 2011).

C Citrus tristeza virus Taxonomic position Genus: Closterovirus

(CTV)

Family: Closteroviridae

Geographical distribution CTV probably originated in Asia and has been disseminated to practically all Citrus spp. growing countries by movement of infected plant material (Balaraman and Ramakrishnan 1977; Manjunath 1985; Valverde et al. 1992; Rahimian 1994; Rocha-Pena et al. 1995; Powell et al. 1997; Cambra et al. 2000; Jarrar et al. 2000; Lee and Bar-Joseph 2000; Dimou et al. 2002; Ahlawat and Pant 2003; Davino et al. 2003; Halbert et al. 2004; Anfoka et al. 2005; Arif et al. 2005; Melzer et al. 2005; Barzegar et al. 2006; Korkmaz et al. 2006; Varveri 2006; Barbarossa et al. 2007; Papayiannis et al. 2007; Biswas 2008; Jiang et al. 2008; Abou Kubaa et al. 2009; Iftikhar et al. 2009; Khan et al. 2009; Albanese et al. 2010; Biswas 2010; Kishore et al. 2010; Sorrentino et al. 2010; Adkar-Purushothama et al. 2011; Cheng et al. 2011; Malandraki et al. 2011; Nelson et al. 2011; Abou Kubaa et al. 2012; Biswas et al. 2012; Chatzivassiliou and Nolasco 2014; Ferretti et al. 2014; Ghosh et al. 2014a; Harper and Pearson 2015; Read and Pietersen 2015; Varveri et al. 2015; Scuderi et al. 2016; Atta et al. 2017; HernandezRodriguez et al. 2017; Read et al. 2017; Singh et al. 2017; Zhou et al. 2017a). Symptoms and host(s) The characteristic symptoms produced by tristeza virus are stunting, stem pitting, chlorosis, and reduced fruit size. Decline of trees occurs on sour orange rootstock (Yang et al. 1999; Cerni et al. 2008). The classic identification procedure for CTV is to graft-inoculate indicator seedlings of Mexican lime (Wallace and Drake 1951) and observe them for vein-clearing, leaf cupping, and stem pitting. There are many strains of CTV and infections frequently occur as mixtures of several strains, it is necessary to be able to distinguish the strains for regulatory purposes, disease management and epidemiology (Miyakawa 1987). Hence techniques were developed to detect CTV and to differentiate the individual strains (Lee et al. 1997; Niblett et al. 2000). The natural hosts of CTV are mostly in the family Rutaceae. The virus infects nearly all citrus species. Some plants in the genus Passiflora are the only hosts of CTV outside the Rutaceae family, apart from the experimental host Nicotiana benthamiana. Transmission The virus is transmitted by a number of aphid vectors: Toxoptera citricidus, Aphis gossypii, A. spiraecola and Toxoptera aurantii; A. gossypii has always been the most efficient vector with transmission efficiencies up to 78%, whereas A. spiraecola and T. aurantii had very low efficiencies of transmission (0–6%) (Raccah et al. 1976; Yokomi and Garnsey 1987; Hermoso de Mendoza et al. 1988; Gottwald et al. 1994; Rocha-Pena et al. 1995; Rahimian et al. 2000; Halbert et al. 2004; Abou Kubaa et al. 2009; Harper et al. 2016). Toxoptera aurantii apparently transmits only certain isolates of CTV. The virus-vector relationship is of a semi-persistent type. The vector acquires and inoculates CTV after feeding probes of 5–16 min. A 24 h acquisition feeding is most effective for virus transmission.

568

Citrus spp. (Citrus aurantifolia; C. aurantium)

Viruliferous aphids lose the ability to transmit after feeding on healthy plants for 24 h or longer. Virus transmissible by mechanical sap-inoculation with low efficiency by stem-slashing methods for experimental purposes (Garnsey et al. 1977); transmissible by grafting; not transmissible by seed. Virion properties and genome The virions are flexuous filaments, with a clear modal length of 2000 nm and 12 nm wide. The genome is a monopartite positive-sense, linear single-stranded RNA of 19,296 nt (NC_001661) which encodes 12 open reading frames that potentially code atleast 17 proteins. Virus coat protein is about 25 kDa (Pappu et al. 1994; Karasev et al. 1995, 1997; Agranovsky and Lesemann 2011).

Citrus variegation virus Taxonomic position Genus: Ilarvirus

(CVV)

Family: Bromoviridae

Geographical distribution CVV infection in plants of Citrus limon was reported by Fawcett and Klotz (1939) from California (USA). The virus occurs in Algeria, Argentina, Australia, India, Israel, Italy, Spain, Uruguay, USA, Lebanon and other citrus growing countries (Fatone et al. 2002; Arif et al. 2005; Loconsole et al. 2009; Abou Kubaa et al. 2015; Rizzo et al. 2017). Symptoms and host(s) Etrog citron and grapefruit usually develop chlorotic leaf symptoms and distortion, which persist on the mature foliage. Infected trees may be stunted and some fruit may be distorted or have chlorotic patterns. Severity differs among isolates. Citrus crinkly leaf is caused by a mild form of CVV. CVV can infect most citrus species viz., lemons, sour orange, citron, and grapefruit on which chlorotic leaf symptoms and distortion occur; mild strains exist that cause only mild mottle or chlorotic flecking. The infected trees are stunted and some fruit may be distorted. Symptoms are reduced in warm weather. Transmission There are no reports of transmission through any type of vectors. The virus is transmissible by mechanical sap-inoculation, and has a broad host range including lemon, sour orange, Etrog citron and grapefruit. Through sap-inoculation the virus is transmitted to herbaceous hosts like some cowpeas (chlorotic or necrotic lesions) and beans (chlorotic mottle or vein-banding). The virus spreads naturally by the distribution of infected planting material. The virus is transmissible by grafting. Certain isolates of CVV are seed-transmitted but at a low incidence. Virion properties and genome The virions are spherical shaped and 26–35 nm in diameter. The genome is tripartite single-stranded plus sense RNA: RNA1 consists of 3433 nt (EF584664 = NC_009537); RNA2 has 2914 nt (NC_009538) and RNA3 has 2309 nt (NC_009536) (Davino and Garnsey 1984; Scott and Ge 1995a; Li et al. 2008; Scott 2011a, b).

Citrus spp. (Citrus aurantifolia; C. aurantium)

Citrus vein enation virus

569

(CVEV)

Synonyms Citrus enation woody gall virus Taxonomic position Genus: Enamovirus

Family: Luteoviridae

Geographical distribution CVEV is also known as woody gall, and was first described in California (Wallace and Drake 1953). The virus-infected Citrus spp. plants were reported from Australia, California, Brazil, India, Japan, Peru, Republic of South Africa, Spain, Turkey, China, and the USA (Mali et al. 1976; Manjunath 1987; da Graca and Maharaj 1991; Vives et al. 2013; Huang et al. 2015; Nakazono-Nagaoka et al. 2015, 2017). Symptoms and host(s) The virus-infected sour orange plants exhibit enations in the leaf veins. The virus also caused woody galls on the trunks or rootstocks of acid lime, rough lemon, rangpur lime and Citrus volkameriana. Transmission The virus is transmitted by aphid vectors Myzus persicae, Toxoptera citricidus and Aphis gossypii (Laird and Weathers 1961; Maharaj and da Graca 1989; Hermoso de Mendoza et al. 1993). The virus is graft-transmissible and infected budwood is the primary source of virus spread. Virion properties and genome The virions are non-enveloped, spherical virion about 25 nm in diameter with T = 3 icosahedral symmetry composed of 180 CP proteins (Maharaj and da Graca 1988; da Graca and Maharaj 1991). The genome is positive-sense single-stranded RNA, with a size of 5,983 nt (HF679486 = NC_021564) coding for five open reading frames (ORFs) (Vives et al. 2013; Huang et al. 2015; Nakazono-Nagaoka et al. 2017).

Citrus viroid V Taxonomic position Genus: Apscaviroid

(CVd V)

Family: Pospiviroidae

Geographical distribution CVd V was first reported in 2007 from Atalantia citroides, a citrus relative. A. citroides propagated on rough lemon (Citrus jambhiri) rootstock (Serra et al. 2008a). The viroid appears to have spread widely in the world (Serra et al. 2008a, b, 2010; Eiras et al. 2009; Cao et al. 2010; Hashemian et al. 2010; Ito and Ohta 2010; Onelge and Yurtmen 2012; Hamdi et al. 2015). Symptoms and host(s) The viroid-infected plants show only very small necrotic lesions and cracks in the stems of the indicator Etrog citron (Citrus medica L.), which sometimes are filled with gum (Verniere et al. 2004).

C

570

Citrus spp. (Citrus aurantifolia; C. aurantium)

Transmission The viroid is mechanically sap-transmissible and also by grafting. Primarily this viroid spreads through the use of infected budwood and root stocks. Etiology and genome properties The genome consists of single-stranded circular RNA. The viroid has a GC-rich genome of 293–294 nts (Serra et al. 2008a); isolates from Pakistan consisted of 292~295 nts (Cao et al. 2013). The 294 nt reference sequence is (EF617306 = NC_010165) (Gora-Sochacka 2004; Giguere et al. 2014).

Citrus viroid VI

(CVd VI)

Synonyms Citrus viroid OS Taxonomic position Genus: Apscaviroid

Family: Pospiviroidae

Geographical distribution CVd VI was first reported in a citrus commercial variety “Shiranui” (Citrus reticulata C. sinensisC. reticulata) grown in Japan (Ito et al. 2001). The viroid spreads in Brazil, China and Japan (Ito et al. 2001, 2002a, b, c; Eiras et al. 2009; Cao et al. 2017a). Symptoms and host(s) The viroid causes mild petiole necrosis and very mild leaf bending in Arizona 861-S1 Etrog citrons (Citrus medica) (Ito et al. 2001). Transmission The viroid is transmitted by grafting, cuttings, and mechanical sap-inoculation. Etiology and genome properties The genome consists of single-stranded circular RNA of 330 nt which is non-coding (AB019508 = NC_004359) (Ito et al. 2001; Gora-Sochacka 2004; Giguere et al. 2014).

Citrus yellow mosaic virus

(CYMV)

Synonyms Citrus mosaic virus Taxonomic position Genus: Badnavirus

Family: Caulimoviridae

Geographical distribution Citrus mosaic disease was first described by Dakshinamurti and Reddy (1975) in South India and subsequently it was renamed as Citrus yellow mosaic virus (CYMV). Ahlawat et al. (1985); Bhaskar Reddy (1997); Sai Gopal et al. 1999; Venkata Prasanna et al. (2002); Ahlawat and Pant (2003); Baranwal et al. 2005; Borah et al. (2009); Rishi (2009); Johnson and Sai Gopal (2012); Johnson

Citrus spp. (Citrus aurantifolia; C. aurantium)

571

et al. (2012); Ghosh et al. (2014b); Anthony Johnson et al. (2017) have reported different aspects of this virus. Symptoms and host(s) The virus induces a bright yellow mottling or vein flecking that persists in mature leaves. Yields are sharply reduced in chronically infected “Sathgudi“ sweet orange trees in India, and fruit may also show mosaic symptoms. The characteristic symptoms of this virus as seen on pommelo, satgudi, and chini sweet are bright yellow mosaic and vein-banding symptoms. The virus can infect most citrus species, cultivars and relatives (Aparna et al. 2002). Transmission The virus is transmitted by the mealybug vector, Planococcus citri in a semi-persistent manner (Pant and Ahlawat 1997; Garnsey et al. 1998). The virus was graft and dodder transmissible to 14 of 15 citrus species or cultivars, and was also transmissible mechanically from citrus to citrus. The disease was also transmitted by mechanical sap-inoculation to, pummelo (6/10), C. decumana (10/10) and Satgudi sweet orange (7/10), but not to grapefruit or Rangpur lime. Infected plants developed typical mosaic symptoms 90 days post-inoculation. Virion properties and genome The virions are bacilliform in shape with a diameter of 30 nm and modal particle length is 130 nm. The genome consists of a single molecule of non-covalently closed circular dsDNA of 7559 bp (NC_003382) (Huang and Hartung 2001). The genome contains three ORFs. The function of protein P1 is unknown, P2 is the virion-associated protein, P3 is a polyprotein with movement protein, coat protein, aspartic protease and RT/RNase H1 domains in that order (Ahlawat et al. 1996a, b; Johnson and Sai Gopal 2012; Johnson et al. 2012; Olszewski and Lockhart 2011; Bhat et al. 2016).

Citrus yellow vein clearing virus Taxonomic position Genus: Mandarivirus

(CYVCV)

Family: Alphaflexiviridae

Geographical distribution Yellow vein disease was first reported from California in the 1950s by Weathers (1960). CYVCV infection in citrus plants was recorded from Pakistan, India, Turkey, Iran and China (Bove 1995; Ahlawat and Pant 2003; Alshami et al. 2003; Onelge et al. 2010, 2011a, 2016; Loconsole et al. 2012a; Chen et al. 2014; Zhen et al. 2015; Cao et al. 2016; Bani Hashmian and Aghajanzadeh 2017; Zhou et al. 2017c). Symptoms and host(s) The virus-infected lemon plants exhibit symptoms of strong yellow vein-clearing, leaf distortion and occasionally ringspots and veinal necrosis. Transmission The virus is mechanically sap-transmissible to mosambi (Citrus limetta) and herbaceous hosts Phaseolus vulgaris, Chenopodium quinoa and C. amaranticolor producing chlorosis, severe mosaic, blotching and necrosis (Alshami et al. 2003). The virus is graft-transmissible. The virus spreads through

C

572

Citrus spp. (Citrus aurantifolia; C. aurantium)

the use of infected propagative material. The virus is not transmissible by seed (Onelge et al. 2011a, b; Zhou et al. 2015). Virion properties and genome The virions are filamentous particles measuring 685 nm in length and 13–14 nm in wide. It has a coat protein of 32 kDa. The genome consists of a positive-sense single-stranded RNA of 7529 nt (JX040635) (Loconsole et al. 2012a), or 7531 nt in the reference sequence (NC_026592) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type, 34 kDa in size (Adams et al. 2004).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Citrus spp. was reported from Turkey and Italy (Paradies et al. 2000). The virus-infected citrus plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Hibiscus green spot virus 2 Taxonomic position Genus: Higrevirus

(HGSV2)

Family: Kitaviridae

HGSV2 infection in plants of Citrus volkameriana was reported from Hawaii (Melzer et al. 2012). The virus is transmitted by mite vectors. For more details of HGSV2, refer to Hibiscus rosa-sinensis.

Hop stunt viroid

(HpSVd)

Synonyms Citrus cachexia viroid; Citrus viroid II Taxonomic position Genus: Hostuviroid

Family: Pospiviroidae

The citrus cachexia disease was first described as a virus-induced, transmissible disease in “Orlando” tangelo (Childs 1950). The viroid nature of cachexia was first suggested in the 1980s (Roistacher et al. 1983). Citrus viroid IIb (CVd-IIb) was identified as the causal agent and named Citrus cachexia viroid (CCaVd) (Semancik et al. 1988). Citrus trees harbor a group of viroids each consists of 295–303 nts called “group II Citrus viroid (or CVd-II)”, which are now classified as the citrus type of Hop stunt viroid (HpSVd-citrus) (Sano et al. 1988). CVd-II (or HpSVd-citrus) is devided into several variants called CVd-IIa, CVd-IIb, CVd-IIc, or Ca-903 by the distinct electrophoretic mobility in polyacrylamide

Citrus spp. (Citrus aurantifolia; C. aurantium)

573

gel; i.e., CVd-IIa (ca. 302–303 nts) is slightly larger than the others (ca. 295–299 nts). Among them, CVd-IIb induced severe gumming, discoloration, and stem-pitting symptoms characteristic of cachexia on the indexing host “Parson’s Special” mandarin, whereas infection by CVd-IIa produced no symptoms on the same host (Semancik et al. 1988). CVd-IIc and Ca-903 also induced symptoms characteristic of cachexia on the indexing host species (Reanwarakorn and Semancik 1999) and CV-IIc was common to xyloporosis disease sources from Israel (Semancik and Duran-Vila 1991), suggesting that cachexia and xyloporosis are the distinct responses of “Orlando” tangero and sweet lime caused by the same agents (Reanwarakorn and Semancik 1999). Cachexia disease and the cachexia-inducing variant of CVd-II cccurs world wide. For example, the CVd-II isolates of HpSVd detected from diseased sweet orange trees (cvs. Navel, Balady andValencia) with gummybark symptoms in Egypt consisted of 299 nt and share 100% identity with CVd-IIb or Ca902 (AF131249; Reanwarakorn and Semancik 1999; Sofy et al. 2010; Hsu et al. 1994; Verniere et al. 2004; Alavi et al. 2006). CVd-II type of HpSVd was also detected from citrus in Jamaica (Bennett et al. 2010). HpSVd infection in Citrus spp. spreads in India, Pakistan, Iran, Israel, Jamaica, Tunisia, Japan, Brazil, Sudan, Taiwan, Syria, Uruguay, Morocco, China, Azerbaijan, Syria and the USA (Diener et al. 1988; Puchta et al. 1989; Hsu et al. 1994; Ito et al. 2002a, b, 2006; Malfitano et al. 2002; Roy and Ramachandran 2003; Ramachandran et al. 2003; Onelge et al. 2004; Palacio-Bielsa et al. 2004; Alavi et al. 2005; Ramachandran et al. 2005; Ragozzino et al. 2005; Elleuch et al. 2006; Kunta et al. 2007; Rizza et al. 2007; Wang et al. 2008a, b, 2010; Mohamed et al. 2009; Abubaker and Elhassan 2010; Bagherian and Izadpanah 2010; Bennett et al. 2010; Sofy et al. 2010; Abou Kubaa et al. 2011; Al-Harthi et al. 2013; Eiras et al. 2013; Pagliano et al. 2013; Mazhar et al. 2014; Sofy and El-Dougdoug 2014; Vamenani et al. 2014; Afechtal et al. 2016; Tan et al. 2016; Najar et al. 2017; Rizzo et al. 2017). The viroid is mechanically sap transmissible. Use of shoots from the infected plants for propagation is the primary cause of this viroid spread. For more details of HpSVd, refer to Humulus lupulus.

Indian citrus ringspot virus Taxonomic position Genus: Mandarivirus

(ICRSV)

Family: Alphaflexiviridae

Geographical distribution ICRSV infection in plants of Citrus spp. was reported from India (Byadgi and Ahlawat 1995; Thind et al. 1995; Pant and Ahlawat 1998; Lore and Cheema 2001; Sharma et al. 2004; Ahlawat and Pant 2003, 2008; Prabha and Baranwal 2012). Four isolates of ICRSV are reported viz., ICRSV-DI, ICRSV-Ab, ICRSV-Ah, and ICRSV-Pu from Delhi, Abohar, Ahmedabad and Pune respectively have been isolated (Hoa and Ahlawat 2004). Symptoms and host(s) The virus-infected citrus plants produces vein-clearing symptoms on young leaves followed by bright yellow rings on mature leaves. Severely affected trees suffer a significant loss of fruit yield and later decline with dieback symptoms. Transmission The virus is mechanically sap-transmissible to Citrus spp. and also to herbaceous hosts including Chenopodium quinoa, C. amaranticolor, soybean, cowpea and French bean cv Saxa in which it became systemic (Pant et al. 1999). The virus is transmitted by bud / grafting, but is not seed-borne or soil-borne

C

574

Citrus spp. (Citrus aurantifolia; C. aurantium)

and has no known vector (Thind et al. 1999; K and Baranwal 2011). It has, however, been detected in pollen. The virus spreads through the use of infected propagative material. Virion properties and genome The virions are flexuous filaments and measure about 650 nm in length and 15 nm width and have helical symmetry with a surface pattern of cross banding. The genome is single-stranded plus sense RNA 7560 nt (AF406744 = NC_003093) and comprises six ORFs, encoding replication related protein, the putative MPs (TGB), the CP and a putative nucleic acid binding regulatory protein. The virus has coat protein of 34 kDa in size (Rustici et al. 2000, 2002; Adams et al. 2004; K and Baranwal 2012).

Iranian citrus ringspot-associated virus

(IrCRSaV)

Taxonomic position IrCRSaV is a tentative member of the genus Cytorhabdovirus and family Rhabdoviridae. Geographical distribution IrCRSaV infection in plants of Citrus spp. was reported from Iran (Sadeghi et al. 2016). Symptoms and host(s) The virus-infected citrus plants exhibit symptoms of small rings, etched lines, and chlorotic spots on both sides of the mature leaves. The rings often followed the lateral veins and turned necrotic as the leaves aged. No specific symptoms were found on fruit. Not all branches of a tree were symptomatic but affected trees were less vigorous than the healthy ones. Transmission The virus is transmissible by mechanical sap-inoculation, and also by grafting. Virion properties and genome The virions are enveloped, bacilliform virions, 30 nm in diameter and 450 nm long. The genome is a monopartite, negative sense, single-stranded RNA. Partial genome sequences are available (KP255975, KP255976) (Sadeghi et al. 2016; Walker et al. 2018).

Olive latent virus 1

(OLV-1)

Taxonomic position Genus: Alphanecrovirus

Family: Tombusviridae

OLV-1 infection in plants of Citrus spp. was reported from Southern Italy (Grieco et al. 1996; Martelli et al. 1996). The virus can infect citrus trees, either exhibiting chlorotic dwarf or no obvious symptoms. The virus is transmitted by fungal vectors and also by mechanical sap-inoculation to herbaceous plants. The virus-infected planting material is the primary source of spread. For more details of OLV-1, refer to Olea europaea.

Citrus spp. (Citrus aurantifolia; C. aurantium)

Satsuma dwarf virus Taxonomic position Genus: Sadwavirus

575

(SDV)

Family: Secoviridae

Geographical distribution SDV was first reported from Japan in Citrus unshia (satsuma) by Yamada and Sawamura (1952). The virus spreads in China, Iran, Korea, Japan, Turkey, Yugoslavia and the former USSR (Ushiyama 1981; Isoda and Gyoutoku 1990; Cui et al. 1991; Iwanami et al. 1991; Zhou et al. 1993). Symptoms and host(s) The virus-infected citrus plants exhibit symptoms of small boat or spoon-shaped leaves. General symptoms are enations, multiple flushing, stunting or dwarfing, reduction in number and size of leaves and shoots, shortened internodes, small-sized fruits with thick peel and reduced fruit production. SDV can infect a wide range of citrus cultivars. All mandarins are susceptible to infection by SDV. Orange, lemon, pummelo, and other hybrids are also susceptible. SDV can infect citrus relatives, including Aegle marmelos, Aeglopsis chevalieri, Atalantia monophylla, Clymenia polyandra, Fortunella polyandra, Citrus trifoliata, and Swinglea glutinosa. Transmission No vector has been identified for this virus. SDV has been mechanically transmitted into various noncitrus hosts such as sesame, Physalis floridana, and herbaceous plants. The virus is transmissible by grafting and use of budwood from infected Citrus spp. is the primary mode of spread of this virus. The virus is not transmissible by contact between plants (Koizumi et al. 1988). Virion properties and genome The virions are non-enveloped 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is bipartite, linear positive-sense, single-stranded RNA. RNA1 consists of 6794 nt (AB009958 = NC_003785) and RNA2 is 5344 nt (AB009959 = NC_003786) (Iwanami et al. 1998, 1999; Iwanami 2008; Sanfacon 2015; Le Gall et al. 2017; Thompson et al. 2017).

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(lemon) in Tucumán Province, Argentina. Eur J Plant Pathol 149:1029–1039 Reanwarakorn K, Semancik JS (1999) Correlation of Hop stunt viroid variants to cachexia and xyloporosis diseases of citrus. Phytopathology 89:568–574 Rishi N (2009) Significant plant virus diseases in India and a glimpse of modern disease management technology. J Gen Plant Pathol 75:1–18 Rizza S, Catara A, Ma X, Deng Z (2007) Detection of multiple infections of Citrus exocortis viroid, Citrus viroid III, and Hop stunt viroid variants in Hunan province, China. Plant Dis 91:1205–1205 Rizzo D, Materazzi A, Stefani L, Panattoni A, Pierro R, De Bellis L, Luvisi A (2017) The occurrence of viruses and viroids in ornamental citrus mother plants in Tuscany (Central Italy). Crop Prot 102:137–140 Rocha-Pena MA, Lee RF, Lastra R, Niblett CL, Ochoa-Corona FM, Garnsey SM, Yokomi RK (1995) Citrus tristeza virus and its aphid vector Toxoptera citricida – threat to citrus production in the Caribbean and central and North America. Plant Dis 79:437–445 Rodrigues JCV, Machado MA, Kitajima EW, Muller GW (2000) Transmission of Citrus leprosis virus by Brevipalpus phoenicis (Acari: Tenuipalpidae). In: da Graca JV, Lee RF, Yokomi RK (eds) Proceedings of the 14th conference of the international organization of citrus virologists. IOCV, Riverside, pp 174–178 Rodrigues JCV, Kitajima EW, Childers CC, Chagas CM (2003) Citrus leprosis virus vectored by Brevipalpus phoenicis (Acari: Tenuipalpidae) on citrus in Brazil. Exp Appl Acarol 30:161–179 Rodrigues JCV, Locali EC, Freitas-Astua J, Kitajima EW (2005) Transmissibility of Citrus leprosis virus by Brevipalpus phoenicis in Solanum violaefolium. Plant Dis 89:911 Rodrigues JCV, Zuniga Reyes JA, Achor DS, Childers CC, Kitajima EW (2007) Occurrence and distribution of Citrus leprosis virus in Honduras. Plant Pathol 56:344 Roistacher CN (1991) Psorosis complex. In: Roistacher CN (ed) Graft-transmissible diseases of Citrus. Handbook for detection and diagnosis. Food and Agriculture Organization of the United Nations, Rome, pp 115–126 Roistacher CN (1993) Psorosis- a review. In: Morenod P, da Graca JV, Timmer LW (eds) Proceedings of the 12th conference of the international organizanization of citrus virologists, IOCV. University of California, Riverside, pp 139–154 Roistacher CN, Gumpf DJ, Nauer EM, Gonzales R (1983) Cachexia disease: virus or viroid. Citrogr 68:111–113 Roistacher CN, Bash JA, Semancik JS (1993) Distinct disease symptoms in Poncirus trifoliate induced by three citrus viroids from three specific groups. In: Moreno P et al (eds) Proceedings of the 12th conference of international organisation of citrus virologists. IOCV, Riverside, pp 173–179 Roman MP, Cambra M, Juarez J, Moreno P, Duran-Vila N, Tanaka FAO, Alves E, Kitajima EW, Yamamoto PT, Bassanezi RB, Teixeira DC, Jesus Junior WC, Ayres AJ, Gimenes-Fernandes N, Rabenstein F, Girotto LF, Bove JM (2004) Sudden death of citrus in Brazil: a graft transmissible bud union disease. Plant Dis 88:453–467 Roy A, Ramachandran P (2003) Occurrence of a Hop stunt viroid (HSVd) variant in yellow corky vein disease of citrus in India. Curr Sci 85:1608–1612 Roy A, Choudhary N, Guillermo LM, Shao J, Govindarajulu A, Achor D, Wei G, Picton DD, Levy L, Nakhla MK, Hartung JS, Brlansky RH (2013) A novel virus of the genus Cilevirus causing symptoms similar to citrus leprosis. Phytopathology 103:488–500 Roy A, Leon MG, Stone AL, Schneider WL, Hartung JS, Brlansky RH (2014) First report of Citrus leprosis virus nuclear type in sweet orange in Colombia. Plant Dis 98:1162 Roy A, Stone AL, Shao J, Otero-Colina G, Wei G, Choudhary N, Achor D, Levy L, Nakhla MK, Hartung JS, Schneider WL (2015a) Identification and molecular characterization of nuclear Citrus leprosis virus, a member of the proposed Dichorhavirus genus infecting multiple Citrus species in Mexico. Phytopathology 105:564–575 Roy A, Leon MG, Stone AL, Schneider WL, Hartung JS, Brlansky RH (2015b) First report of Citrus leprosis virus nuclear type in sweet orange in Colombia. J Virol Methods 224:105–109 Roy A, Hartung JS, Schneider WL, Shao J, León MG, Melzer MJ, Otero-Colina G, Beard JJ, Bauchan GR, Ochoa R, Brlansky RH (2015c) Role bending: complex relationships between viruses, hosts and mite vectors related to citrus leprosis, an emerging disease. Phytopathology 105:1013–1025

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586

Citrus spp. (Citrus aurantifolia; C. aurantium)

Rustici G, Accotto GP, Noris E, Masenga V, Luisoni E, Milne RG (2000) Indian citrus ringspot virus: a proposed new species with some affinities to potex-, carla-, fovea- and allexiviruses. Arch Virol 145(9):1895–1908 Rustici G, Milne RG, Accotto GP (2002) Nucleotide sequence, genome organisation and phylogenetic analysis of Indian citrus ringspot virus. Arch Virol 147(11):2215–2224 Saavedra FS, Bernal A, Childers C, Kitajima E (2001) First report of citrus leprosis on Panama. Plant Dis 85(2):228 Sadeghi MS, Afsharifar A, Izadpanah K, Loconsole G, De Stradis A, Martelli GP, Saponari M (2016) Isolation and partial characterization of a novel Cytorhabdovirus from citrus trees showing foliar symptoms in Iran. Plant Dis 100:66–71 Sai Gopal DVR, Sreenivasula M, Raghavendra Rao G, Ventakaprasana TM and Subbaiah KV (1999) Characterization and identification of a Citrus mosaic badnavirus infecting acid lime (Citrus aurantifolia) in Andhra Pradesh. pp 845–852. In: Hitech citrus management. Proceedings of the international sympossium on citriculture. Nagpur Sanchez de la Torre ME, Riva O, Zandomeni R, Grau O, Garcia ML (1998) The top component of Citrus psorosis virus contains two ssRNAs, the smaller encodes the coat protein. Mol Plant Pathol Online. http://www.bspp.org.uk/mppol/ 1998/1019sanchez Sanchez de la Torre ME, Lopez C, Grau O, Garcia ML (2002) RNA 2 of Citrus psorosis virus is of negative polarity and has a open reading frame in its complimentary strand. J Gen Virol 83:1777–1781 Sanfacon H (2015) Secoviridae: A family of plant picorna-like viruses with monopartite or bipartite genomes. In: Encyclopedia of Life Sciences. Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000764.pub3 http:// www.els.net Sano T, Hataya T, Shikata E (1988) Complete nucleotide sequence of a viroid isolated from Etrog citron, a new member of Hop stunt viroid group. Nucleic Acids Res 16:347 Scott SW (2011a) Bromoviridae and Allies. In: Encyclopedia of Life Sciences (ELS). Wiley, Chichester. https://doi.org/ 10.1002/9780470015902 Scott SW (2011b) Ilarvirus. Bromovwiridae. In: The springer index of viruses. Springer, New York, pp 187–194. https:// doi.org/10.1007/978-0-387-95919-1_27 Scott SW, Ge X (1995a) The complete nucleotide sequence of RNA-3 of Citrus leaf rugose and citrus variegation Ilarviruses. J Gen Virol 76:957–963 Scott SW, Ge X (1995b) The nucleotide sequence of Citrus leaf rugose virus RNA-1. J Gen Virol 76:3233–3238 Scuderi G, Russo M, Davino S, Ferraro R, Catara A, Licciardello G (2016) Occurrence of the T36 genotype of Citrus tristeza virus in citrus orchards in Sicily, Italy. Plant Dis 100:1253 Semancik JS (1988) Citrus exocortis viroid – a review. In: Proceedings of the 10th conference of ICOV. IOCV, Riverside, pp 136–151 Semancik JS, Duran-Vila N (1991) The grouping of citrus viroids: Additional physical and biological determinants and relationships with diseases of citrus. In: Proceedings of the 11th conference of the international organization of citrus virologists. IOCV, Riverside, pp 178–187 Semancik JS, Vidalakis G (2005) The question of Citrus viroid IV as a Cocadviroid. Arch Virol 150:1059–1067 Semancik JS, Weathers LG (1972) Exocortis virus: an infectious free nucleic acid plant virus with unusual properties. Virology 47:456–466 Semancik JS, Roistacher CN, Rivera-Bustamante R, Duran-Vila N (1988) Citrus cachexia viroid, a new viroid of citrus: relationship to viroids of the exocortis disease complex. J Gen Virol 69:3059–3068 Semancik JS, Szychowski JA, Rakowski AG, Symons RH (1993) Isolates of Citrus exocortis viroid recovered by host and tissue selection. J Gen Virol 74:2427–2436 Semancik JS, Rakowski AG, Bash JA, Gumpf DJ (1997) Application of selected viroids for dwarfing and enhancement of production of “Valencia” orange. J Hortic Sci 72:563–570 Serra P, Barbosa CJ, Daros JA, Flores R, Duran-Vila N (2008a) Citrus viroid V: molecular characterization and synergistic interactions with other members of the genus Apscaviroid. Virology 370:102–112 Serra P, Eiras M, Bani-Hashemian SM, Murcia N, Kitajima EW, Daros JA, Flores R, Duran-Vila N (2008b) Citrus viroid V: occurrence, host range, diagnosis, and identification of new variants. Phytopathology 98:1199–1204 Serra P, Barbosa CJ, Daros JA, Flores R, Duran-Vila N (2010) Identification and characterization of a variant of Citrus viroid V (CVd-V) in Seminole Tangelo. In: Proceedings of the 17th conference of the international organization of citrus virologists, Riverside, California, vol 17. IOCV, Riverside, pp 150–157 Sharma S, Singh B, Rani G, Nagpai A, Zaidi AA, Hallan V, Virk GS (2004) Current status of Indian citrus ringspot virus (ICRSV) in Kinnow orchards of Punjab and neibouring states. Int J Biosci Rep 2(2):132–135 Singh AK, Meetai NT, Singh BK, Mandal N (2017) High incidence of Citrus tristeza virus in mandarin (Citrus reticulata) in north-east states of India. Virus Dis 28:401–407 Sofy AR, El-Dougdoug KA (2014) First record of a Hop stunt viroid variant associated with gumming and stem pitting on Citrus volkameriana trunk rootstock in Egypt. New Dis Rep 30:11

Citrus spp. (Citrus aurantifolia; C. aurantium)

587

Sofy AR, Soliman AM, Mousa AA, Ghazal SA, El-Dougdoug KA (2010) First record of Citrus viroid II (CVd-II) associated with gummy bark disease in sweet orange (Citrus sinensis) in Egypt. New Dis Rep 21:24 Sorrentino G, Davino S, Davino M (2010) Distrribution of three different isolates of Citrus tristeza virus in southern Italy. J Plant Pathol 92:S4.101 Tan SH, Talibov THO, Krueger RR, Bodaghi S, Dang T, Chao YY, Greer G, Vidalakis G (2016) First report of Citrus exocortis viroid and two citrus variants of the Hop stunt viroid on lemon in Azerbaijan. Plant Dis 100:2341 Tanner JD, Kunta M, da Graca JV, Skaria M, Nelson SD (2010) Evidence for citrus tatter leaf seed transmission in citrus. Citrus research and technology: XVIII conference of the IOCV (Campinas (SP), Brazil), 31:50 Tessitori M, La Rosa R, Catara A (2002) Citrus psorosis virus bark scaling on tarocco sweet orange. Plant Dis 86:560 Thind SK, Kapur SP, Sharma JN (1995) Citrus ring spot virus – a new record from Punjab. Plant Dis Res 10(1):75–77 Thind SK, Arora PK, Sharma JN, Cheema SS (1999) Transmission of Citrus ring spot virus through budwood, seed and insect-vectors. Indian J Virol 15(1):47–48 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J GenVirol 98:529–531 Ushiyama K (1981) Studies on satsuma dwarf disease. II. Spread of the disease by topworking satsuma trees with infected scions. Bulletin of the Kanagawa Horticultural Experiment Station No. 28, 24–30 Valverde RA, Bourgeois J, Lee RF (1992) First report of Citrus tristeza virus in Louisiana. Plant Dis 80:103 Vamenani R, Rahimian H, Alavi SM, Babaeizad V (2014) Detection of Hop stunt viroid in lemon, sweet lime, Clementine and Satsuma mandarins and grapefruit trees in Mazandaran. Iran J Plant Pathol 50(2):Pe191–Pe191, En105 Varveri C (2006) Molecular detection and characterization of Citrus tristeza virus in Greece. Phytopathol Mediterr 45:68 Varveri C, Olmos A, Pina JA, Marroquin C, Cambra M (2015) Biological and molecular characterization of a distinct Citrus tristeza virus isolate originating from a lemon tree in Greece. Plant Pathol 64(4):792–798 Venkata Prasanna TM, Subbaiah KV, Seshadri KV, Sai Gopal DVR (2002) Survey and indexing of yellow mosaic virus disease on sweet orange (Citrus sinensis) and acid lime (Citrus aurantifolia) in Andhra Pradesh. Indian J Citricult 1:83–86 Verniere C, Perrier X, Dubois C, Dubois A, Botella L, Chabrier C, Bove JM, Duran Vila N (2004) Citrus viroids: symptom expression and effect on vegetative growth and yield of clementine trees grafted on trifoliate orange. Plant Dis 88:1189–1197 Villalobos W, Rivera C, Hammond RW (1997) Occurrence of citrus viroids in Costa Rica. Rev Biol Trop 45:983–987 Visvader JE, Gould AR, Bruening GE, Symons RH (1982) Citrus exocortis viroid: nucleotide sequence and secondary structure of an Australian isolate. FEBS Lett 137:288–292 Vives MC, Galipienso L, Navarro L, Moreno P, Guerri J (2001) The nucleotide sequence and genomic organization of Citrus leaf blotch virus: candidate type species for a new virus genus. Virology 287:225–233 Vives MC, Galitienso L, Navarro L, Moreno P, Guerri J (2002) Citrus leaf blotch virus: a new citrus virus associated with bud union crease on trifoliate root stocks. In: Duran-Vila N, Milne RG, Da Graca JV (eds) Proceedings of the 15th conference of International Organization of Citrus Virologists. IOCV, Riverside, pp 205–212 Vives MC, Moreno P, Navarro L, Guerri J (2008a) Citrus leaf blotch virus. In: Rao GP, Myrta A, Ling K (eds) Characterization, diagnosis and management of plant viruses, vol 2. Studium Press, Houston, pp 55–67 Vives MC, Martìn S, Ambrus S, Renovell A, Navarro L, Pina JA, Moreno P, Guerri J (2008b) Development of a fullgenome cDNA clone of Citrus leaf blotch virus and infection of citrus plants. Mol Plant Pathol 9:787–797 Vives MC, Velazquez K, Pina JA, Moreno P, Guerri J, Navarro L (2013) Identification of a new Enamovirus associated with citrus vein enation disease by deep sequencing of small RNAs. Phytopathology 103:1077–1086 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Rhabdoviridae. J GenVirol 99:447–448 Wallace JM, Drake RJ (1951) Recent developments in studies of quick decline and related diseases. Phytopathology 41:785–793 Wallace JM, Drake RJ (1953) A virus-induced vein enation in citrus. Citrus Leaves 33:22–24 Wallace JM, Drake RJ (1962) Tatterleaf, a previously undescribed virus effect of citrus. Plant Dis Reptr 46:211–212 Wang XF, Zhou CY, Tang KZ, Lan JQ, Zhou Y, Li ZA (2008a) Preliminary studies on species and distribution of citrus viroids in China. Agric Sci China 7:1097–1103 Wang XF, Zhou CY, Tang KZ, Li ZA (2008b) Occurrence of four citrus viroids in Chongqing, China. Plant Dis 92:978–978 Wang XF, Zhou Y, Li ZG, Tang KZ, Liu YQ, Cao MJ, Zhou CY (2010) Molecular, biological and phylogenetic analysis of Chinese isolates of Hop stunt viroid associated with citrus cachexia disease. J Phytopathol 158(5):372–377

C

588

Cladrastis kentukea (Kentucky yellowwood)

Weathers LG (1960) Yellow-vein disease of citrus and studies on interactions between yellow-vein and other viruses of citrus. Virology 11:753–764 Yamada S, Sawamura K (1952) Studies on the dwarf disease of Satsuma orange, Citrus unshiu Markov. (preliminary report). Hortic Div TokaiKinki Agr Exp Sta Bull 1:61–67 Yang ZN, Mathews DM, Dodds JA, Mirkov TE (1999) Molecular characterization of an isolate of Citrus tristeza virus that causes severe symptoms in sweet orange. Virus Genes 19:131–142 Yokomi RK, Garnsey SM (1987) Transmission of Citrus tristeza virus by Aphis gossypii and Aphis citricola in Florida. Phytophylactica 19:169–172 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J GenVirol 98:131–133 Zhang TM, Liang XY, Roistacher CN (1988) Occurrence and detection of Citrus tatter leaf virus (CTLV) in Huangyan, Zhejiang Province, China. Plant Dis 72:543–545 Zhen S, Kurth EG, Peremyslov VV, Changyong Z, Dolja VV (2015) Molecular characterization of a Citrus yellow vein clearing virus strain from China. Arch Virol 160:1811–1813 Zhou C, Zhao X, Jiang Y, He X (1993) The occurrence of Satsuma dwarf virus in China. In: Moreno P, da Graca JV, Timmer LW (eds) Proceedings of the 12th conference of the International Organisation of Citrus Virologists, pp 349–351 Zhou Y, Chen HM, Wang XF, Li ZA, Tang M, Zhou CY (2015) Lack of evidence for seed transmission of Citrus yellow vein clearing virus despite its frequent detection in seed tissues. J Plant Pathol 97(3):1–3 Zhou Y, Liu Y, Liu K, Yang F, Zhou C (2017a) Distribution and population structure of Citrus tristeza virus in Poncirus trifoliata. Australas Plant Pathol 46:351–355 Zhou Y, Zhang Y, Liu Y, Chen H, Li T, Zhou C (2017b) Distribution and molecular characterization of Citrus chlorotic dwarf-associated virus in China. Australas Plant Pathol 46:227–229 Zhou Y, Chen HM, Cao MJ, Wang XF, Jin X, Liu KH, Zhou CY (2017c) Occurrence, distribution and molecular characterization of Citrus yellow vein clearing virus in China. Plant Dis 101:137–143

Cladrastis kentukea (Kentucky yellowwood) Synonyms Cladrastis tinctoria; C. lutea Family: Fabaceae

Ornamental

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Cladrastis lutea was reported from New York (Provvidenti and Hunter 1975). The virus-infected Kentucky yellowwood plants exhibit symptoms of veinal chlorosis, mottle, rolling along the main veins, and necrosis, and plant growth was retarded. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

References Provvidenti R, Hunter JE (1975) Bean yellow mosaic virus infection in Cladrastis lutea, an ornamental leguminous tree. Plant Dis Reptr 59:86–87

Claytonia virginica (Spring beauty)

589

Clarkia amoena (Godetia) Synonyms Godetia grandiflora Family: Onagraceae

C Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Clarkia amoena/Godetia grandiflora was reported from Italy (Roggero et al. 1999). The virus-infected godetia plants show necrotic ring symptoms. The virus is transmitted by a thrips vector, Frankliniella occidentalis, in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Clarkia amoena was reported by Cho et al. (1987) and Parrella et al. (2003). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Cho JJ, Mau RFL, Mitchell WC, Gonsalves D, Yudin LS (1987) Host list of plants susceptible to Tomato spotted wilt virus (TSWV). University of Hawaii, Honolulu, 10 p Parrella G, Gognalons P, Gebre-Selassi K, Vovlas C, Marchoux G (2003) An update of the host range of Tomato spotted wilt virus. J Plant Pathol 85(4, special issue):227–264 Roggero P, Ciuffo M, Dellavalle G, Gotta P, Gallo S, Peters D (1999) Additional ornamental species as hosts of Impatiens necrotic spot tospovirus in Italy. Plant Dis 83:967

Claytonia virginica (Spring beauty) Family: Montiaceae

Medicinal

Spring beauty latent virus Taxonomic position Genus: Bromovirus

(SBLV)

Family: Bromoviridae

590

Clematis spp.

Geographical distribution SBLV infection was first reported in Claytonia virginica plants from Arkansas, USA (Valverde 1985). The virus spreads in the USA (Valverde and Glascock 1991). Symptoms and host(s) The virus-infected spring beauty plants exhibit symptomless infections. Transmission The virus is transmitted by beetle vectors. The virus is also transmissible by mechanical sap-inoculation, and by grafting. The virus is not transmissible by seed. Virion properties and genome The virions are non-enveloped about 28 nm in diameter with T = 3 icosahedral symmetry, composed of 180 capsid proteins: 12 pentamers and 20 hexamers. The genome consists of three linear ssRNA components. The complete genome sequence has been generated from the infectious clone of SBLV, which revealed that the RNA-1 genome segment is 3252 nt (AB080598), RNA-2 is 2898 nt (AB080599), and RNA-3 is 2213 nt (AB080600) (Valverde 1985; Valverde and Glascock 1991; Fujisaki et al. 2003; Kao and Adkins 2011; Scott 2011).

References Fujisaki K, Hagihara F, Kaido M, Mise K, Okuno T (2003) Complete nucleotide sequence of pring beauty latent virus, a bromovirus infectious to Arabidopsis thaliana. Arch Virol 148(1):165–175 Kao CC, Adkins S (2011) Bromovirus. Bromoviridae. In: The Springer index of Viruses. Springer, New York, pp 173–177. https://doi.org/10.1007/978-0-387-95919-1_25 Scott SW (2011) Bromoviridae and allies. In: Encyclopedia of Life Sciences (ELS). Wiley, Chichester. https://doi.org/ 10.1002/9780470015902 Valverde RA (1985) Spring beauty latent bromovirus: a new member of the Bromovirus group. Phytopathology 75:395–398 Valverde RA, Glascock CB (1991) Further examination of the RNA and coat protein of Spring beauty latent virus. Phytopathology 81:401–404

Clematis spp. Family: Ranunculaceae

Ornamental

Apple mosaic virus

(ApMV)

Taxonomic position Genus: Ilarvirus

Family: Bromoviridae

ApMV infection in plants of Clematis vitalba was reported from Turkey (Arli Sokmen et al. 2005; Mitrofanova et al. 2018). The virus-infected clematis plants were symptomless. No insect vector is known for this virus. The virus is mechanically sap-transmissible. The virus spreads by the use of vegetative planting material from the infected plants, and virus is graft-transmissible. For more details of ApMV, refer to Malus domestica.

Clematis spp.

591

Clematis chlorotic mottle virus Taxonomic position Genus: Pelarspovirus

(ClCMoV)

Family: Tombusviridae

Geographical distribution ClCMoV infection in plants of Clematis spp. was reported from the USA and Russia (Mollov et al. 2014; McLaughlin et al. 2017; Mitrofanova et al. 2018; Zakubanskiy et al. 2018). Symptoms and host(s) Several varieties of Clematis from domestic and international sources show symptoms of yellow mottling and veining, chlorotic ringspots, line pattern mosaics, and in some cases flower distortion and discoloration. Transmission There is no known vector for this virus. The virus is transmitted by mechanical sap-inoculation. The virus spread is through the use of infected vegetative propagative material. Virion properties and genome The virions are isometric and icosahedral (T = 3), not enveloped, 28–34 nm in diameter. The genome is monopartite, linear single stranded positive sense RNA of 3880 nt (KX712140 = NC_033777) with five open reading frames (ORFs) encoding a 27-kDa protein (ORF 1), an 87-kDa replicase protein (ORF 2), two centrally located movement proteins (ORF 3 and 4), and a 37-kDa capsid protein (ORF 5) (McLaughlin et al. 2017).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Clematis spp. was reported from the UK and New Zealand (Dingley 1969; Guy 2011; Mitrofanova et al. 2018). The virus-infected clematis plants exhibit leaf mottle and chlorotic vein-banding symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation Clematis spp. to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Plum pox virus Taxonomic position Genus: Potyvirus

(PPV)

Family: Potyviridae

PPV infection in plants of Clematis spp. was reported from Slovenia (Virscek Marn et al. 2004). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PPV, refer to Prunus domestica.

C

592

Clematis spp.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Clematis vitalba was reported from Yugoslavia, Italy, and the UK (Bellardi et al. 1985; Rana et al. 1987; Harju et al. 2011; Mitrofanova et al. 2018). The virus-infected clematis plants exhibit chlorotic spots or yellow vein netting symptoms. The virus is transmitted by the thrips vectors, is present in/on the pollen, and enters into the host through the injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Clematis paniculata plants was reported from New Zealand (Thomson 1978). The virus-infected clematis plants exhibit chlorotic spots, rings, and ringspot lesions, with some necrosis and leaf distortion symptoms. The virus is transmitted by nematode vectors in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and also by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Clematis afoliata was reported from New Zealand (Dingley 1969; Mitrofanova et al. 2018). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Arli Sokmen M, Kutluk Yilmaz ND, Mennan H, Sevik MA (2005) Natural weed hosts of Apple mosaic virus in hazelnut orchards in Turkey. J Plant Pathol 87:239–242 Bellardi MG, Credi R, Gelli C (1985) Tobacco streak virus in Clematis vitalba L. Phytopathol Mediterr 24:255–259 Dingley JM (1969) Records of plant diseases in New Zealand. NZ Dep Sci Ind Res Bull 192:1–298 Guy PL (2011) Detection of Cucumber mosaic virus on Clematis paniculata in lowland forest in New Zealand. Aust Plant Dis Notes 6:20–21 Harju VA, Skelton AL, Monger WA, Forde SMD, Daly M, Nixon T, Lawson R, Bennett S, Mumford RA (2011) A report of viruses, viroids, and phytoplasmas found in ornamental plants from 1999–2007: findings from Central Science Laboratory, UK. Acta Hortic 901:223–229 McLaughlin M, Lockhart B, Jordan R, Denton G, Mollov D (2017) Complete nucleotide sequence of Clematis chlorotic mottle virus, a new member of the family Tombusviridae. Arch Virol 162:1373–1379 Mitrofanova IV, Zakubanskiy AV, Mitrofanova OV (2018) Viruses infecting main ornamental plants: an overview. Ornamental Hortic 24:95–102

Cleome gynandra (Cat’s whiskers)

593

Mollov D, Lockhart B, Phibbs A, Creswell T, Ruhl G, Dorman E, Kinard G, Jordan R (2014) Clematis chlorotic mottle virus, a novel virus occurring in clematis in the USA. Phytopathology 104(S3):81 Rana GL, Krajačić M, Štefanac Z, Pleše N, Rubino L, Miličić D (1987) Properties of a new strain of Tobacco streak virus from Clematis vitalba (Ranunculaceae). Ann Appl Biol 111:153–160 Thomson AD (1978) An unidentified virus-like disease of Clematis paniculata. N Z J Bot 16:167–168 Virscek Marn M, Mavric I, Urbancic-Zemljic M, Skerlavaj V (2004) Detection of Plum pox virus Potyvirus in weeds. Acta Hortic 657:251–254 Zakubanskiy A, Mitrofanova I, Chirkov S (2018) First report of Clematis chlorotic mottle virus on clematis in Russia. J Plant Pathol 100(3):605–605

Cleome gynandra (Cat’s whiskers) Family: Capparaceae

Weed host

Ageratum enation virus Taxonomic position Genus: Begomovirus

(AEV)

Family: Geminiviridae

AEV infection in plants of Cleome gynandra was reported from India (Raj et al. 2010). The virusinfected Cat’s whiskers plants exhibit leaf curl symptoms including upward curling of the leaves and crinkling, shortening of petioles, swelling of nodes and internodes, and stunting of the whole plant. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of AEV, refer to Ageratum spp.

Cleome golden mosaic virus Taxonomic position Genus: Begomovirus

(CleGMV)

Family: Geminiviridae

Geographical distribution CleGMV infection in plants of Cleome gynandra was reported from Brazil (Fonseca et al. unpublished HQ396465). Symptoms and host(s) The virus-infected cleome plants exhibit golden mosaic symptoms. Transmission Insect transmission of CleGMV has not been demonstrated. However, in common with all other begomoviruses, CleGMV is likely transmitted by the whitefly vector of begomoviruses Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The virion structure of ClGMV has not been investigated. In common with all geminiviruses, the virions of ClGMV are likely geminate (twinned quasi-icosahedra).

C

594

Cleome gynandra (Cat’s whiskers)

It is unclear at this time whether CleGMV is a bipartite Begomovirus, as is typical of the majority of begomoviruses native to the New World, or monopartite since only the sequence of a single genome/ DNA-A genomic component of 2566 nt (HQ396465 = NC_015397) is available (Brown et al. 2015; Zerbini et al. 2017). Typical of the DNA-A component of bipartite begomoviruses and the genomes of monopartite begomoviruses native to the New World, the sequence is predicted to encode five genes, one in the virion-sense and four in the complementary-sense. The expression and function of these genes have not been investigated for CleGMV.

Cleome leaf crumple virus Taxonomic position Genus: Begomovirus

(CleLCrV)

Family: Geminiviridae

Geographical distribution CleLCrV infection in plants of Cleome gynandra was reported from Brazil (da Silva et al. 2011; Wyant et al. 2012). Symptoms and host(s) The virus-infected cleome plants exhibit leaf leaf crumpling, yellow mosaic and stunting symptoms. Transmission Insect tranmsision of CleLCrV has not been demonstrated. However, in common with all other begomoviruses, CleLCrV is likely transmitted by the whitefly vector of begomoviruses Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The virion structure of CleLCrV has not been investigated. In common with all geminiviruses, the virions of CleLCrV are likely geminate (twinned quasi-icosahedra). CleLCrV is typical of the majority of begomoviruses native to the New World with a bipartite genome consisting of two circular, single-stranded DNA components. DNA-A contains 2725 nt (JF694461 = NC_016578; FN435999) and DNA-B of 2661 nt (JF694462 = NC_016572; FN436000) (Briddon 2001; Paprotka et al. 2010; Wyant et al. 2012; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the New World, the DNA-A component of CleLCrV encodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for CleLCrV.

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 da Silva S, Castillo-Urquiza G, Hora Júnior B, Assunção I, Lima G, Pio-Ribeiro G, Mizubuti E, Zerbini F (2011) High genetic variability and recombination in a Begomovirus population infecting the ubiquitous weed Cleome affinis in northeastern Brazil. Arch Virol 156:2205–2213

Clerodendrum spp.

595

Paprotka T, Metzler V, Jeske H (2010) The first DNA 1-like alpha satellites in association with New World begomoviruses in natural infections. Virology 404:148–157 Raj SK, Snehi SK, Khan MS, Tiwari AK, Rao GP (2010) Detection of Ageratum enation virus from cat’s whiskers (Cleome gynandra L.) with leaf curl symptoms in India. J Gen Plant Pathol 76:292–294 Wyant PS, Strohmeier S, Schafer B, Krenz B, Assuncao IP, Lima GS, Jeske H (2012) Circular DNA genomics (circomics) exemplified for geminiviruses in bean crops and weeds of northeastern Brazil. Virology 427(2):151–157 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Clerodendrum spp. Synonyms Clerodendron spp. Family: Lamiaceae

Medicinal

Clerodendrum chlorotic spot dichorhavirus Taxonomic position Genus: Dichorhavirus

(ClCSV)

Family: Rhabdoviridae

Geographical distribution ClCSV infection in plants of Clerodendrum spp. was reported from Brazil and Central Amazon basin (Kubo et al. 2007, 2008; Kitajima et al. 2003, 2008; Rodrigues et al. 2008; Takassuigui 2009; Takassuigui et al. 2010; Ramos-González et al. 2018). Symptoms and host(s) The virus-infected clerodendrum plants exhibit chlorotic and necrotic spots on leaves, or dark green spots and ringspots on their leaves. Transmission The virus is transmitted by mite vectors, Brevipalpus phoenicis and B. yothersi. The virus is also mechanically sap-transmissible (Kitajima et al. 2008; Kubo et al. 2008; Ramos-González et al. 2018). Virion properties and genome The virions are short bacilliform and measures 40 nm  100–110 nm in size. The genome is composed of two single-stranded negative-sense RNAs. RNA1 consists of 6524 nt (MG938506) and RNA2 is 6105 nt (MG938507); RNA1 includes five ORFs, and RNA2 has a single ORF encoding the RdRp (Ramos-González et al. 2018; Walker et al. 2018).

Clerodendron golden mosaic virus Taxonomic position Genus: Begomovirus

(ClGMV)

Family: Geminiviridae

C

596

Clerodendrum spp.

Geographical distribution ClGMV infection in plants of Clerodendrum philippinum was reported from Vietnam and China (Ha et al. 2008; Ding et al. 2010, 2012). Symptoms and host(s) The virus-infected clerodendrum plants exhibit chlorosis and yellow mosaic symptoms. Transmission Insect transmission of ClGMV has not been demonstrated. However, in common with all other begomoviruses, ClGMV is likely transmitted by the whitefly vector of begomovirus Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The virion structure of ClGMV has not been investigated. In common with all geminiviruses, the virions of ClGMV are likely geminate (twinned quasi-icosahedra). ClGMV is typical of the majority of begomoviruses native to the New World with a bipartite genome consisting of two circular, single-stranded DNA components. DNA-A contains 2767 nt (DQ641692 = NC_010713) and DNA-B 2757 nt (DQ641693 = NC_010714) (Briddon 2001; Ha et al. 2008; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the New World, the DNA-A component of ClGMV encodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for ClGMV.

Clerodendron yellow mosaic virus Taxonomic position Genus: Begomovirus

(ClYMV)

Family: Geminiviridae

Geographical distribution ClYMV infection in plants of Clerodendrum inerme was reported from New Delhi (India) (Sivalingam et al. 2011). Symptoms and host(s) The virus-infected clerodendrum plants exhibit bright chlorotic spots along the midrib, reduction in leaf lamina, leaf deformation, and stunting. The virus has been isolated from Clerodendrum inerme with chlorosis and yellow mosaic symptoms, Codiaeum variegatum with mild leaf curling and foliar yellowing, and Bougainvillea peruviana with leaf curling and stunting symptoms. Transmission The transmission of ClYMV has not been investigated. It is likely that, in common with other begomoviruses, ClYMV is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of ClYMV has not been investigated. In common with all geminiviruses, the virions of ClYMV are likely geminate (twinned quasi-icosahedra).

Clerodendrum spp.

597

ClYMV is a typical Old World monopartite Begomovirus with a genome consisting of a single circular molecule of single-stranded DNA of 2760 nt (EF408037 = NC_009451; HE863667, KF704391) (Briddon 2001; Sivalingam et al. 2011; Brown et al. 2015; Zerbini et al. 2017). The characterized genome of ClYMV encodes the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated.

Clerodendrum golden mosaic China virus Taxonomic position Genus: Begomovirus

(ClGMCNV)

Family: Geminiviridae

Geographical distribution ClGMCNV infection in plants of Clerodendrum cyrtophyllum was reported from Jiangsu Province, China, and the USA (Li and Zhou 2010; Ding et al. 2012; Valverde et al. 2012). Symptoms and host(s) The virus-infected clerodendrum plants exhibit yellow mosaic, yellow vein, and mild leaf distortion symptoms. ClGMCNV has been identified infecting Clerodendrum cyrtophyllum and Salvia splendens with foliar yellow mosaic symptoms. Transmission The transmission of the ClGMCNV isolate from China has not been investigated. It is likely that, in common with other begomoviruses, ClGMCNV is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. Attempts to transmit the US isolate using Bemisia tabaci were unsuccessful, although the virus was shown to be graft-transmissible (Valverde et al. 2012). Experimentally the virus originating from China has been introduced into Nicotiana benthamiana (inducing leaf crumpling and stunting), Nicotiana glutinosa (inducing chlorotic spots and leaf distortion), and Nicotiana tabacum (inducing leaf distortion, stunting) plants by Agrobacterium-mediated inoculation of clones. Infections of Petunia hybrida were symptomless. Virion properties and genome The virion structure of ClGMCNV has not been investigated. In common with all geminiviruses, the virions of ClGMCNV are likely geminate (twinned quasi-icosahedra). The genome of ClGMCNV is bipartite consisting of two circular, single-stranded DNA components. DNA-A contains 2776 nt (FJ011668 = NC_011346; FN396962) and DNA-B of 2739 nt (FJ011669 = NC_011347; FN396963) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the Old World, the DNA-A component of ClGMCNV encodes six genes, two in the virion-sense and four in the complementary-sense, and the DNA-B encodes two genes, one in each orientation (Li and Zhou 2010). The expression and function of these genes have not been investigated for ClGMCNV.

Clerodendrum golden mosaic Jiangsu virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

(ClGMJsV)

C

598

Clerodendrum spp.

Geographical distribution ClGMJsV infection in plants of Clerodendrum cyrtophyllum was reported from China (Li and Zhou 2010). Symptoms and host(s) The virus-infected clerodendrum plants exhibit foliar yellow mosaic symptoms. Transmission The transmission of ClGMJsV has not been investigated. It is likely that, in common with other begomoviruses, ClGMJsV is transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Experimentally the virus has been introduced into Nicotiana benthamiana (inducing downward leaf curling), Nicotiana glutinosa (inducing mild downward leaf curling and slight stunting), Nicotiana tabacum (inducing mild downward leaf curling and slight stunting), and Petunia hybrida (inducing upward leaf curling, vein swelling, and stunting) plants by Agrobacterium-mediated inoculation of clones (Li and Zhou 2010). Virion properties and genome The virion structure of ClGMJsV has not been investigated. In common with all geminiviruses, the virions of ClGMJsV are likely geminate (twinned quasi-icosahedra). ClGMJsV is a typical Old World monopartite Begomovirus. Only a single isolate of the virus has been characterized, and the genome consists of a single circular molecule of single-stranded DNA of 2753 nt (FN396966) (Briddon 2001; Li and Zhou 2010; Brown et al. 2015; Zerbini et al. 2017). The characterized genome of ClGMJsV encodes the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Clerodendrum thomsoniae was reported from Wisconsin (Khan and Maxwell 1975). The virus-infected clerodendrum plants show characteristic ringspot symptoms consisting of chlorotic yellow rings, necrotic rings, puckering, chlorosis, and distortion of the leaves. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sapinoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Yam mosaic virus Taxonomic position Genus: Potyvirus

(YMV)

Family: Potyviridae

Clerodendrum spp.

599

YMV infection in plants of Clerodendrum umbellatum was reported from Lome, Togo (West Africa), by Gumedzoe (1993). The virus-infected clerodendrum plants exhibit mosaic mottling symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of YMV, refer to Dioscorea spp.

C References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Ding M, Wang Y, Fang Q, Zhang ZK (2010) Occurrence of Clerodendrum golden mosaic virus in Clerodendrum bungei in Yunnan China. J Plant Pathol 92:S4.120 Ding M, Wang Y, Fang Q, Zhang ZK (2012) Occurrence of Clerodendrum golden mosaic virus in Clerodendrum bungei in Yunnan China. J Plant Pathol 92:S4.107–S4.122 Gumedzoe MY (1993) Major virus diseases of medicinal plants in West Africa. Acta Hortic (ISHS) 331:307–310 Ha C, Coombs S, Revill P, Harding R, Vu M, Dale J (2008) Molecular characterization of begomoviruses and DNA satellites from Vietnam: additional evidence that the New World geminiviruses were present in the Old World prior to continental separation. J Gen Virol 89:312–326 Khan MA, Maxwell DP (1975) Identification of Tobacco ring spot virus in Clerodendrum thomsoniae. Phytopathology 65:1150–1153 Kitajima EW, Chagas CM, Rodrigues JCV (2003) Brevipalpus-transmitted plant virus and virus-like diseases: cytopathology and some recent cases. Exp Appl Acarol 30:135–160 Kitajima EW, Kubo KS, Ferreira PTO, Alcantara BK, Boari AJ, Gomes RT, Freitas-Astua J, Rezende JAM, Morais GJ, Salaroli RB (2008) Chlorotic spots on Clerodendrum, a disease caused by a nuclear type of Brevipalpus (Acari: Tenuipalpidae) transmitted virus. Sci Agric 65:36–49 Kubo KS, Antonioli-Luizon R, Freitas-Astua J, Kitajima EW (2007) Molecular diagnosis of the Clerodendron chlorotic spot virus (ClCSV). Summa Phytopathol 33(supplement):S12 Kubo KS, Freitas-Astúa J, Kitajima EW (2008) Detection of Clerodendrum chlorotic spot virus in plants and its mite vector. Summa Phytopathol 34(Suppl):S97 Li J, Zhou X (2010) Molecular characterization and experimental host-range of two begomoviruses infecting Clerodendrum cyrtophyllum in China. Virus Genes 41(2):250–259 Ramos-González PL, Chabi-Jesus C, Banguela-Castillo A, Tassi AD, Rodrigues MDC, Kitajima EW, Harakava R, Freitas-Astúa J (2018) Unveiling the complete genome sequence of clerodendrumchlorotic spot virus, a putative Dichorhavirus infecting ornamental plants. Arch Virol 163(9):2519–2524 Rodrigues JCV, Antony LMK, Salaroli RB, Kitajima EW (2008) Brevipalpus-associated viruses in the central Amazon basin. Trop Plant Pathol 33:12–19 Sivalingam PN, Satheesh V, John P, Chandramohan S, Malathi VG (2011) Complete genome sequence of an isolate of Clerodendron yellow mosaic virus – a new Begomovirus from India. Acta Virol 55(4):357–360 Takassuigui GR (2009) Biological and molecular characterization of Clerodendrum chlorotic spot virus. Master thesis Takassuigui GR, Kitajima EW, Tanaka FAO, Marques JPR, Appezzato-da-Gloria B (2010) Anatomy of lesions caused by Clerodendrum chlorotic spot virus (ClCLV) transmitted by the mite Brevipalpus phoenicis in different host species. Summa Phytopathol 36:291–297 Valverde RA, Sabanadzovic S, Hammond J (2012) Viruses that enhance the aesthetics of some ornamental plants: beauty or beast? Plant Dis 96:600–611 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Rhabdoviridae. J GenVirol 99:447–448 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

600

Clitoria ternatea (Butterfly pea)

Clitoria ternatea (Butterfly pea) Family: Fabaceae

Medicinal

Clitoria chlorosis virus

(ClCV)

Taxonomic position ClCV is a tentative member of the genus Potyvirus and family Potyviridae Geographical distribution ClCV infection in plants of Clitoria ternatea was reported from Western Australia (Coutts et al. 2011). Symptoms and host(s) The virus-infected Clitoria ternatea plants showed chlorosis, without other obvious symptoms. Transmission The virus is presumed to be transmitted by aphid vectors. Virion properties and genome The virions are flexuous filaments. The genome consists of a single molecule of positive-sense singlestranded RNA, presumed to be of c.9.5–10 kb. A partial genome sequence of 840 nt is available (JF427593) (Coutts et al. 2011; Wylie et al. 2017).

Clitoria virus Y Taxonomic position Genus: Potyvirus

(ClVY)

Family: Potyviridae

Geographical distribution ClVY infection in plants of Clitoria ternatea was reported from Australia (Mackenzie et al. unpublished - AF228515). Symptoms and host(s) The virus-infected butterfly pea plants exhibit mosaic symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner, and the virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense, single-stranded RNA. A partial genome sequence of 1720 nt is available (AF228515) (Revers and Garcia 2015; Wylie et al. 2017).

Clitoria ternatea (Butterfly pea)

601

Clitoria yellow mottle virus Taxonomic position Genus: Tobamovirus

(CliYMV)

Family: Virgaviridae

Geographical distribution CliYMV infection in plants of Clitoria ternatea was reported from Northern Australia (Wei et al. 2012). Symptoms and host(s) The virus-infected butterfly pea seedlings show local chlorotic lesions and a systemic vein-clearing followed by mottling and mosaic of the tip leaves. Transmission There is no known vector for this virus. The virus is mechanically sap-transmissible to a number of host plants. The virus is also transmissible by contact between plants. Virion properties and genome The virions are rigid rod shaped of 18 nm in diameter and have a predominant length of 300–310 nm. The genome consists of a single molecule of linear positive-sense single-stranded RNA of 6514 nt JN566124 = NC_016519 (Wei et al. 2012). The 30 terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G50 pppG) (Zaitlin 2011; Adams et al. 2017).

Clitoria yellow vein virus Taxonomic position Genus: Tymovirus

(CYVV)

Family: Tymoviridae

Geographical distribution CYVV was first reported in plants of Clitoria ternatea from Kenya by Bock et al. (1977). The virus spreads in Kenya. Symptoms and host(s) The virus-infected butterfly pea plants exhibit systemic vein chlorosis symptoms. Transmission The virus is transmitted by beetle vectors in a semi-persistent manner. The virus is also transmissible by mechanical sap-inoculation. The virus is not transmitted by the seed. Virion properties and genome The virions are non-enveloped, isometric capsid with T = 3 icosahedral symmetry, about 30 nm in diameter. The genome consists of a single molecule of positive-sense ssRNA. The 30 terminus has a tRNA-like structure. Partial genome sequences are available (AF035200; M15963) (Martelli et al. 2002).

C

602

Cnidoscolus acontifolius (Chaya)

Cowpea chlorotic mottle virus Taxonomic position Genus: Bromovirus

(CCMV)

Family: Bromoviridae

CCMV infection in plants of Clitoria ternatea was reported from Nigeria (Thottappilly et al. 1993). The virus-infected butterfly pea plants exhibit bright yellow mosaic symptoms, which varied from small circular spots or stipples to large patches, occasionally covering the entire leaflet. The virus is transmitted by beetle vectors and also by mechanical sap-inoculation. For more details of CCMV, refer to Vigna unguiculata.

Peanut stunt virus Taxonomic position Genus: Cucumovirus

(PSV)

Family: Bromoviridae

PSV infection in plants of Clitoria ternatea was reported from Sudan (Ahmed and Mills 1985). The virus-infected butterfly pea plants exhibit green-yellow mottle symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PSV, refer to Arachis hypogaea.

References Adams MJ, Adkins S, Bragard C, Gilmer D, Li D, MacFarlane SA, Wong S-M, Melcher U, Ratti C, Ryu KH, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Virgaviridae. J GenVirol 98:1999–2000 Ahmed AH, Mills PR (1985) Identification of Peanut stunt virus in the Sudan. Plant Dis 69:173–174 Bock KR, Guthrie EJ, Meredith G (1977) Clitoria yellow vein virus, a Tymovirus from Kenya. Ann Appl Biol 85:97–103 Coutts BA, Kehoe MA, Webster CG, Wylie SJ, Jones RA (2011) Indigenous and introduced potyviruses of legumes and Passiflora spp. from Australia: biological properties and comparison of coat protein nucleotide sequences. Arch Virol 156(10):1757–1774 Martelli GP, Sabanadzovic S, Abou-Ghanem Sabanadzovic N, Edwards MC, Dreher T (2002) The family Tymoviridae. Arch Virol 147(9):1837–1846 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Thottappilly G, Sehgal OP, Rossel HW (1993) Characteristics of a Cowpea chlorotic mottle virus isolate from Nigeria. Plant Dis 77:60–63 Wei K, Gibbs A, Mackenzie A (2012) Clitoria yellow mottle virus: a tobamovirus from Northern Australia. Australas Plant Dis Notes 7:59–61 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J GenVirol 98:352–354 Zaitlin M (2011) Tobamovirus. In: Tidona C, Darai G (eds) The Springer index of viruses. Springer, New York. https://doi. org/10.1007/978-0-387-95919-1

Cnidoscolus acontifolius (Chaya) Synonyms Cnidoscolus chayamansa Family: Euphorbiaceae

Leafy vegetable

Cnidoscolus urens (Bull Nettle)

603

Cassava common mosaic virus Taxonomic position Genus: Potexvirus

(CsCMV)

Family: Alphaflexiviridae

CsCMV infection in plants of Cnidoscolus chayamansa was reported from Venezuela, Tuvalu, and Mexico (Elliott and Zettler 1987; Jones et al. 1998; Mejias et al. 2015). The virus-infected chaya plants show extensive mosaic and leaf deformation symptoms. The virus is transmitted by means not involving a vector. Primarily the virus spread takes place through the use of infected cuttings for planting. The virus is transmissible by mechanical sap-inoculation. For more details of CsCMV, refer to Manihot esculenta.

References Elliott MS, Zettler FW (1987) Cassava common mosaic virus infections of chaya (Cnidoscolus aconitifolius) in Yucatan, Mexico. Plant Dis 71:353–356 Jones P, Devonshire J, Dabek A, Howells C (1998) First report of Cassava common mosaic potexvirus infecting Chaya (Cnidoscolus chayamansa) in Tuvalu. Plant Dis 82:591 Mejias A, Rodriguez-Roman E, Romano M, Zambrano K, Marys E (2015) New record of Cassava common mosaic virus infecting Chaya (Cnidoscolus chayamansa McVaug) in Venezuela. Plant Dis 99:1190

Cnidoscolus urens (Bull Nettle) Family: Euphorbiaceae

Weed host

Cnidoscolus mosaic leaf deformation virus Taxonomic position Genus: Begomovirus

(CnMLDV)

Family: Geminiviridae

Geographical distribution CnMLDV infection in plants of Cnidoscolus urens was reported from Brazil (Melo et al. 2016). Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative nonpropagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2641 nt (KT966771 = NC_038982) and DNA-B of 2613 nt (KT966772) (Briddon 2001; Brown et al. 2015; Melo et al. 2016; Zerbini et al. 2017).

C

604

Coccinia spp. (C. barteri, C. cordifolia, C. grandis, and C. indica) (Ivy gourd)

References Briddon RW (2001) Begomovirus. Geminiviridae. The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Melo AM, Silva SJ, Ramos-Sobrinho R, Ferro MM, Assunção IP, Lima GS (2016) Cnidoscolus mosaic leaf deformation virus: a novel Begomovirus infecting euphorbiaceous plants in Brazil. Arch Virol 161:2605–2608 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Coccinia spp. (C. barteri, C. cordifolia, C. grandis, and C. indica) (Ivy gourd) Family: Cucurbitaceae

Vegetable

Coccinia mosaic Tamil Nadu virus Taxonomic position Genus: Begomovirus

(CMTNV)

Family: Geminiviridae

Geographical distribution CMTNV infection in plants of Coccinia grandis was reported from India (Nagendran et al. 2016). Symptoms and host(s) The virus-infected Ivy gourd plants exhibit mosaic symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2749 nt (KM244719 = NC_024810) and DNA-B of 2691 nt (KM244718 = NC_024809) (Briddon 2001; Brown et al. 2015; Nagendran et al. 2016; Zerbini et al. 2017).

Coccinia mottle virus Taxonomic position Genus: Ipomovirus

(CocMoV)

Family: Potyviridae

Coccinia spp. (C. barteri, C. cordifolia, C. grandis, and C. indica) (Ivy gourd)

605

Geographical distribution CocMoV infection in plants of Coccinia grandis was reported from Sudan (Desbiez et al. 2016). Symptoms and host(s) The virus infected ivy gourd plants exhibit mottling symptoms. The virus has a narrow natural host range within the cucurbits, and a low prevalence. Transmission The virus is transmitted by the Mediterranean biotype of whitefly vector Bemisia tabaci in a nonpersistent manner, but not by the aphid Myzus persicae. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long and 11–13 nm wide. The genome contains single molecule of linear, positive sense single-stranded RNA of 9778 nt in length (KU935732 = NC_030840) (Desbiez et al. 2016; Wylie et al. 2017).

Moroccan watermelon mosaic virus Taxonomic position Genus: Potyvirus

(MWMV)

Family: Potyviridae

MWMV infection in plants of Coccinia barteri was reported from Southeastern Nigeria (Owolabi et al. 2012). The virus-infected ivy gourd plants exhibit symptoms of vein-clearing and sometimes yellow mosaic, green vein-banding which sometimes may be masked by white blotches on the leaves. The virus is transmitted by aphid vectors, Myzus persicae and Aphis gossypii, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of MWMV, refer to Citrullus lanatus.

Papaya ringspot virus Taxonomic position Family: Potyviridae

(PRSV)

Genus: Potyvirus

PRSV infection in plants of Coccinia spp. was reported from Tamilnadu (India) (Nagendran et al. 2017). The virus is spread by a number of different aphid species including the green peach and melon aphids in a non-persistent manner. The virus is also mechanically sap-transmissible. For more details of PRSV, refer to Carica papaya.

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

C

606

Cocos nucifera (Coconut palm)

ToLCNDV infection in plants of Coccinia indica was reported from Northern India (Raj et al. 2012). The virus-infected ivy gourd plants exhibit yellow mosaic and curling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is graft-transmissible. For more details of ToLCNDV, refer to Solanum lycopersicum.

References Briddon RW (2001) Begomovirus. Geminiviridae. The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Desbiez C, Verdin E, Tepfer M, Wipf-Scheibel C, Millot P, Dafalla G, Lecoq H (2016) Characterization of a new cucurbitinfecting Ipomovirus from Sudan. Arch Virol 161(10):2913–2915 Nagendran K, Satya VK, Mohankumar S, Karthikeyan G (2016) Molecular characterization of a distinct bipartite Begomovirus species infecting ivy gourd (Cocciniagrandis L.) in Tamil Nadu, India. Virus Genes 52:146–151 Nagendran K, Mohankumar S, Aravintharaj R, Balaji CG, Manoranjitham SK, Singh AK, Rai AB, Singh B, Karthikeyan G (2017) The occurrence and distribution of major viruses infecting cucurbits in Tamil Nadu state, India. Crop Prot 99:10–16 Owolabi AT, Rabenstein F, Ehrig F, Maiss Edgar M, Vetten HJ (2012) Strains of Moroccan watermelon mosaic virus isolated from Lagenaria breviflorus and Coccinia barteri in Calabar, Southeastern Nigeria. Int J Virol 8:258–270 Raj SK, Snehi SK, Khan MS, Singh R, Tiwari AK, Rao GP (2012) Biological, molecular studies and management of begomoviruses affecting cucurbitaceous crops in India. Recent trends in plant virology. Govind P Rao, V K Baranwal, Bikash Mandal, Narayan Rishi. Studium Press LLC, Houston 135–155 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J GenVirol 98:352–354 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Cocos nucifera (Coconut palm) Family: Arecaceae

Oil seed

Coconut cadang-cadang viroid Taxonomic position Genus: Cocadviroid

(CCCVd)

Family: Pospiviroidae

Geographical distribution “Cadang-cadang” disease of Cocos nucifera has been known in the Philippines since 1914 (Hanold and Randles 1989, 1991a, b, 1998), and a disease with the symptoms now known to be specific for cadangcadang was first described on San Miguel Island in 1931 (Ocfemia 1931). The viroid spreads in the Philippines, Guam, and Malaysia (Price 1971; Randles 1975; Zelazny 1979; Zelazny and Niven 1980; Zelazny et al. 1982; Randles et al. 1980, 1991; Mohammadi et al. 2013).

Cocos nucifera (Coconut palm)

607

Symptoms and host(s) The earliest symptoms in naturally infected coconut palm are rounding of nut shape; the development of equatorial nut scarifications; and the appearance of fine, translucent, bright yellow leaf spots. Inflorescences then become necrotic, nut production declines and then ceases, frond production slows, and general chlorosis appears, followed by death of the crown. Eight to 16 years elapse between first symptoms and death of the palm. Artificially inoculated seedlings show varying severities of leaf spotting and stunting. Some palms die soon, those that continue to develop never flower. Estimated total losses exceed 40 million coconut palms. CCCVd natural host range includes Cocos nucifera L., Elaeis guineensis Jacquin, and Corypha elata Roxburgh (Randles et al. 1980, 1998; Rodriguez 1993). The experimental host range of CCCVd includes certain species in the Arecaceae, such as Adonidia merrillii (Manila palm), Areca catechu (betel nut palm), Caryota cumingii (fishtail palm), Chrysalidocarpus lutescens (yellow butterfly palm), Livistona rotundifolia (round-leaf fountain palm), Phoenix dactylifera (date plum), Ptychosperma macarthurii (Macarthur palm), and Roystonea regia (royal palm) (Imperial et al. 1985).

Transmission In the field, natural transmission is observed but the mechanism remains as yet unknown. A low rate of seed transmission has been observed, and pollen transmission is suspected (Pacumbaba et al. 1994), but these results need to be confirmed. Artificial inoculation has been achieved by a high-pressure injection of nucleic acid extracts into the shoots of germinating nuts. Experimental hosts susceptible to mechanical sap-inoculation with partially purified CCCVd include a wide range of Arecaceae including oil palm, betel nut palm (Areca catechu), date palm, royal palm, and golden cane palm (Imperial et al. 1985). Like other viroid diseases, there is a possibility of CCCVd transmission via contaminated cutting tools or farm equipment, even transmission from articles made of coconut materials cannot be excluded.

Etiology and genome properties The genome consists of a single-stranded circular RNA. CCCVd is the smallest known viroid, as well as the smallest known infectious pathogen (246-nucleotide form). Cadang-cadang-affected coconut palms (Cocos nucifera) contain linear and circular viroid molecules, and there are numerous variants of the basic 246 nucleotide sequence, i.e., 246, 247, 287, and 297 nts (J02049) (Haseloff et al. 1982; Rodriguez and Randles 1993; Gora-Sochacka 2004; Vadalamai et al. 2008; Flores et al. 2011; Giguere et al. 2014). The reference sequence is 246 nt (NC_001462).

Coconut foliar decay virus Taxonomic position Genus: Unassigned

(CFDV)

Family: Nanoviridae

Geographical distribution CFDV infection was first reported in plants of Cocos nucifera from Saraoutou, Vanuatu, by Calvez et al. (1980). The virus spreads in Vanuatu (Julia 1982; Randles et al. 1987).

C

608

Cocos nucifera (Coconut palm)

Symptoms and host(s) The typical initial symptoms are that a frond between positions 5 and 11 below the spear leaf which first shows chlorosis in leaflets, over about one quarter of the frond. The whole frond becomes chlorotic, as do the immediately adjacent fronds, so that a central whorl in the crown appears yellow. These fronds collapse when marginal necrosis is observed. They hang down through the normal lower whorls of the crown. The severity of symptoms depends on the cultivar, and in some cases symptoms disappear in a “remission” phase. In susceptible coconut palms like the Malayan Red Dwarf, the crown dies within 6 months to 2 years after symptoms first appear. Transmission The virus is transmitted by the planthopper vector, Myndus taffini Bonfils (Cixiidae), in a semipersistent manner or in a persistent manner (Wefels et al. 2015). No seed or mechanical transmission has been demonstrated. This virus is also not transmissible by grafting (Julia 1982; Julia et al. 1985). Virion properties and genome It is an icosahedral virus, 20 nm in diameter, with an associated positive-sense circular single-stranded DNA of 1291 nt (M29963 = NC_001465) is now described as an alphasatellite) (Randles et al. 1987, 1992; Randles and Hanold 1989; Rohde et al. 1990). Three circular single-stranded DNAs of 1286 nt (DNA S.1, MF926436), 1263 nt (DNA S.2, MF926439), and 641 nt (DNA gamma, MF926442) are associated with isolate VU-89 (Gronenborn et al. 2018).

Coconut tinangaja viroid Taxonomic position Genus: Cocadviroid

(CTiVd)

Family: Pospiviroidae

Geographical distribution Tinangaja disease was first reported as a destructive disease of Cocos nucifera on Guam in 1917 (Boccardo 1985). The viroid is widespread in the Guam islands (Rodriguez and Carpio 1987; Hodgson et al. 1998; Randles et al. 1998). Symptoms and host(s) The viroid-infected coconut palms show a progressive thinning of the crown, reduction in the size and number of fruit, shriveling and deformation of fruit, cessation of fruit production, sterile inflorescences, and total sterility and may take up to 15 years to die (Wall and Randles 2003). Transmission The viroid is successfully transmitted mechanically by a high-pressure injection (Wall and Randles 2003). There is some evidence that CTiVd can infect pollen grains, although pollen transmission has not yet been demonstrated. Etiology and genome properties The genome consists of a single-stranded circular RNA of 254 nt, which is noncoding (M20731 = NC_001471) (Keese et al. 1988; Gora-Sochacka 2004; Giguere et al. 2014).

Cocos nucifera (Coconut palm)

609

References Boccardo G (1985) Viroid aetiology of tinangaja and its relationship with cadang-cadang disease of coconut. In: Maramorosch K, MacKelvey JJ (eds) Subviral pathogens of plants and animals: viroids and prions. Academic, Orlando, pp 75–99 Calvez C, Renard JL, Marty G (1980) Tolerance of the hybrid coconut local x Rennell to new Hebrides disease. Oleagineux 35:443–451 Flores R, Daros J, Hernandez J-A, Di Serio F (2011) Viroids. In: Encyclopedia of life sciences (ELS). John Wiley and Sons Ltd, Chichester. https://doi.org/10.1002/9780470015902.a0000434.pub3 Giguere T, Raj Adkar-Purushothama C, Perreault J-P (2014) Comprehensive secondary structure elucidation of four genera of the family Pospiviroidae. PLoS One 9(6):e98655. https://doi.org/10.1371/journal.pone.0098655 Gora-Sochacka A (2004) Viroids: unusual small pathogenic RNAs. Acta Biochim Pol 51(3):587–607 Gronenborn B, Randles JW, Knierim D, Barriere Q, Vetten HJ, Warthmann N, Cornu D, Sileye T, Winter S, Timchenko T (2018) Analysis of DNAs associated with coconut foliar decay disease implicates a unique single-stranded DNA virus representing a new taxon. Sci rep 8(1):5698 Hanold D, Randles JW (1989) Cadang-cadang-like viroid in oil palm in the Solomon Islands. Plant Dis 73(2):183 Hanold D, Randles JW (1991a) Coconut Cadang-cadang disease and its viroid agent. Plant Dis 75(4):330–335 Hanold D, Randles JW (1991b) Detection of coconut cadang cadang viroid-like sequences in oil and coconut palm and other monocotyledons in the south-west Pacific. Ann Appl Biol 118:139–151 Hanold D, Randles JW (1998) Report on ACIAR-funded research on viroids and viruses of coconut palm and other tropical monocotyledons 1985–1993. ACIAR monograph, no. 45. Australian Centre for International Agricultural Research, Canberra, 222 pp Haseloff J, Mohamed NA, Symons RH (1982) Viroid RNAs of cadang-cadang disease of coconuts. Nature, UK 299:316–321 Hodgson RAJ, Wall GC, Randles JW (1998) Specific identification of Coconut tinangaja viroid for differential field diagnosis of viroids in coconut palm. Phytopathology 88:774–781 Imperial JS, Bautista RM, Randles JW (1985) Transmission of the Coconut cadang-cadang viroid to six species of palm by inoculation with nucleic acid extracts. Plant Pathol 34(3):391–401 Julia JF (1982) Myndus taffini (Homoptera cixiidae), vector of foliar decay of coconuts in Vanuatu. Oleagineux 37:409–414 Julia JF, Dollet M, Randles JW, Calvez C (1985) Foliar decay of coconut by Myndus taffini (FDMT). Oleagineux 40:19–27 Keese P, Osorio-Keese ME, Symons RH (1988) Coconut tinangaja viroid: sequence homology with Coconut cadangcadang viroid and other Potato spindle tuber viroid related RNAs. Virology 162:508–510 Mohammadi M-R, Meon S, Wong M-Y, Daros J-A, Vadamalai G (2013) Characterization of variants of Coconut cadangcadang viroid from coconut palm in Malaysia. International Workshop on Viroids and Satellite RNAs (IWVds) Beijing, 23–25 Aug, p 20 Ocfemia GO (1931) The probable nature of cadang-cadang disease of coconuts. Philipp Agric 26:338–340 Pacumbaba EP, Zelazny B, Orense JC, Rillo EP (1994) Evidence for pollen and seed transmission of Coconut cadangcadang viroid in Cocos nucifera. J Phytopathol 142:37–42 Price WC (1971) Cadang-cadang of coconut – a review. Plant Sci 3:1–13 Randles JW (1975) Association of two ribonucleic acid species with cadang-cadang disease of coconut palm. Phytopathology 65:163–167 Randles JW, Hanold D (1989) Coconut foliar decay virus particles are 20-nm icosahedra. Intervirology 30:177–180 Randles JW, Boccardo G, Imperial JS (1980) Detection of the cadang-cadang associated RNA in African oil palm and buri palm. Phytopathology 70(3):185–189 Randles JW, Hanold D, Julia JF (1987) Small circular single-stranded DNA associated with foliar decay disease of coconut palm in Vanuatu. J Gen Virol 68:273–280 Randles JW, Rodriguez MJB, Imperial JS (1988) Cadang-cadang disease of coconut palm. Microbiol Sci 5:18–22 Randles JW, Hanold D, Pacumbaba EP, Rodriguez MJB (1991) Cadang-cadang disease of coconut palm. In: Mukhopadhyay AN, Kumar J, Chaube HS, Singh US (eds) Plant diseases of international importance. Prentice Hall, Englewood Cliffs Randles JW, Miller DC, Morin JP, Rohde W, Hanold D (1992) Localisation of Coconut foliar decay virus in coconut palm. Ann Appl Biol 121:601–617 Randles JW, Hanold D, Pacumbaba EP, Rodriguez MJB (1998) Cadang-cadang disease of coconut palm – an overview. In: Randles JW, Hanold D (eds) Report on ACIAR-funded research on viroids and viruses of coconut palm and other tropical monocotyledons 1985–1993. ACIAR working paper no. 51, Aug 1998. ISBN 1-86320-219-6 Rodriguez MJB (1993) Molecular variations in Coconut cadang-cadang viroid (CCCVd). PhD Thesis. University of Adelaide, South Australia Rodriguez MJB, Carpio CB (1987) Survey report on the tinangaja disease and coconut varieties/populations in Guam, USA. 21 pp

C

610

Codiaeum variegatum (Garden croton)

Rodriguez MJB, Randles JW (1993) Coconut cadang-cadang viroid (CCCVd) mutants associated with severe disease vary in both the pathogenicity domain and the central conserved region. Nucleic Acids Res 21:2771 Rohde W, Randles JW, Langridge P, Hanold D (1990) Nucleotide sequence of a circular single-stranded DNA associated with Coconut foliar decay virus. Virology 176:648–651 Vadalamai G, Perera AAFLK, Hanold D, Rezaian MA, Randles JW (2008) Detection of Coconut cadang-cadang viroid sequences in oil palm and coconut palm by ribonuclease protection assay. Ann Appl Biol 154:117–125 Wall GC, Randles JW (2003) Coconut tinangaja viroid. In: Hadidi A et al (eds) Viroids, vol 36. CSIRO Publishing, Collingwood, pp 242–245, Part IV Wefels E, Morin JP, Randles JW (2015) Molecular evidence for a persistent-circulative association between Coconut foliar decay virus and its vector Myndus taffini. Australas Plant Pathol 44:283–288 Zelazny B (1979) Distribution and spread of the cadang-cadang disease of coconut palm. Acta Phytopathol Acad Sci Hung 14:115 Zelazny B, Niven BS (1980) Duration of the stages of cadang-cadang disease of coconut palm. Plant Dis 64:841–842 Zelazny B, Randles JW, Boccardo G, Imperial JS (1982) The viroid nature of the Cadang-cadang disease of coconut palm. Sci Filipinas 2:46–63

Codiaeum variegatum (Garden croton) Family: Euphorbiaceae

Ornamental

Clerodendron yellow mosaic virus Taxonomic position Genus: Begomovirus

(ClYMV)

Family: Geminiviridae

ClYMV infection in plants of Codiaeum variegatum was reported from Lahore (Pakistan) (Anwar et al. 2012). The virus-infected garden croton plants show mild leaf curl and yellowing symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ClYMV, refer to Clerodendrum spp.

Eggplant mottled dwarf nucleorhabdovirus

(EMDV)

Synonyms Croton vein yellowing virus Taxonomic position Genus: Nucleorhabdovirus

Family: Rhabdoviridae

EMDV infection in plants of Codiaeum variegatum was reported from Italy (Bellardi et al. 1991; Bertaccini and Bellardi 1992; Parrella et al. 2012). The virus-infected garden croton plants exhibit yellow and/or pink vein-banding and stunting symptoms. The virus is transmitted by the leafhopper vectors in a persistent, propagative manner. The virus is mechanically sap-transmissible and also by grafting. For more details of EMDV, refer to Solanum melongena.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(INSV)

Codonanthe spp.

611

INSV was reported to infect Codiaeum variegatum in Iran (Ghotbi and Shahraeen 2012; Ghotbi 2013). The virus-infected garden croton plants exhibit symptoms of necrotic leaf spot, chlorosis, and stunting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV was reported to infect Codiaeum variegatum in Iran (Ghotbi and Shahraeen 2012). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Anwar S, Tahir M, Zaidi NSS, Briddon RW (2012) First report of Clerodendron yellow mosaic virus infecting croton. J Plant Pathol 94:S4.101 Bellardi MG, Bertaccini A, Villani A, Vicci V (1991) First report of Rhabdovirus like particles in croton (Codiaeum variegatum). Plant Dis 75:862 Bertaccini A, Bellardi MG (1992) A Rhabdovirus inducing vein yellowing in croton. Plant Pathol 41:79–82 Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iran J Plant Pathol 49:41 Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses affecting propagated ornamentals in Northern Iran. Int Res J Microbiol 3:373–381 Parrella G, Greco B, De Stradis A, Cavicchi L, Bellardi MG (2012) Serological and molecular evidence that Croton vein yellowing virus is an isolate of Eggplant mottled dwarf virus. J Plant Pathol 94(Supp 4):S4.73

Codonanthe spp. Family: Gesneriaceae

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

Ornamental

(TMV)

Family: Virgaviridae

TMV was reported to infect cultivated Codonanthe spp. in the USA (Zettler and Nagel 1983). The symptoms on the foliage of infected codonanthe plants were inconspicuous. There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Reference Zettler FW, Nagel J (1983) Infection of cultivated gesneriads by two strains of Tobacco mosaic virus. Plant Dis 67:1123–1125

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612

Coffea arabica (Coffee)

Coffea arabica (Coffee) Family: Rubiaceae

Commercial crop

Coffee ringspot dichorhavirus Taxonomic position Genus: Dichorhavirus

(CoRSV)

Family: Rhabdoviridae

Geographical distribution CoRSV infection in plants of Coffea arabica was first reported from the State of Sao Paulo by Bitancourt (1938). The virus spreads in Brazil, Costa Rica, and the Philippines (Rodrigues et al. 2002; Chagas et al. 2003; Kitajima and Chagas 2009; Kitajima et al. 2011; Ramalho et al. 2014, 2016). Symptoms and host(s) Coffee plants infected with this virus display conspicuous localized ringspot lesions on both leaves and berries. The disease is capable of causing significant leaf and fruit drop with accompanying reduced coffee berry yields. Transmission The virus is transmitted by a mite vector, Brevipalpus phoenicis, in a persistent propagative manner (Chagas 1973; Chagas et al. 2003). The virus is transmissible by mechanical sap-inoculation, and produces necrotic local lesions on Chenopodium spp. (Chagas et al. 1981). The virus is seed-transmitted in Coffea excelsa to the extent of 8.1% (Reyes 1961). Virion properties and genome The virions are bacilliform and non-enveloped and measure 40  100–110 nm. The genome is comprised of two molecules of negative-sense single-stranded RNA: RNA1 consists of 6552 nt (KF812525) and RNA2 of 5945 nt (KF812526) (Kitajima and Costa 1972; Chagas 1980; Boari et al. 2004; Locali et al. 2005; Dietzgen et al. 2014; Ramalho et al. 2014; Walker et al. 2018).

References Bitancourt AA (1938) Coffee ring spot, a new disease of coffee. O Biol 4:404–405 Boari AJ, Freitas-Astua J, Ferreira PTO, Neder DG, Nogueira NL, Rossi ML, Kitajima EW (2004) Purification and serology of the Coffee ringspot virus. Summa Phytopathol 30:453–458 Chagas CM (1973) Association of Brevipalpus phoenicis (Geijskes) to the coffee ring spot. O Biol 39:229–232 Chagas CM (1980) Morphology and intracellular behavior of Coffee ringspot virus (CRV) in tissues of coffee (Coffea arabica L.). Phytopathol Z 99:301–309 Chagas CM, July JR, Alba APC (1981) Mechanical transmission and structural features of Coffee ring spot virus (CRV). Phyto Pathologisches Z 102:100–106 Chagas CM, Kitajima EW, Rodrigues JCV (2003) Coffee ring spot virus vectored by Brevipalpus phoenicis (Acari: Tenuipalpidae) in coffee. Exp Appl Acarol 30:203–213 Dietzgen RG, Kuhn JH, Clawson AN, Freitas-Astua J, Goodin MM, Kitajima EW, Kondo H, Wetzel T, Whitfield AE (2014) Dichorhavirus: a proposed new genus for Brevipalpus mite-transmitted, nuclear, bacilliform, bipartite, negative-strand RNA plant viruses. Arch Virol 159:607–619 Kitajima EW, Chagas CM (2009) Viral diseases in coffee. In: Wintgens JN (ed) Coffee: growing, processing, sustainable production, 2nd edn. Wiley-VCH, Weinheim, pp 550–556 976p Kitajima EW, Costa AS (1972) Bacilliform particles associated to the coffee ring spot. Cienc Cult 24:542–545

Coix chinensis (Coix seed)

613

Kitajima EW, Chagas CM, Braghini MT, Fazuoli LC, Locali-Fabris EC, Salaroli RB (2011) Natural infection of several Coffea species and hybrids and Psilanthus ebracteolatus by the Coffee ringspot virus (CoRSV). Sci Agric (Piracicaba, Braz) 68:503–507 Locali EC, Freits-Astua J, Antonioli-Lauizon R, Boari AJ, Machado MA (2005) Diagnosis of the coffee ringspot by RT-PCR. Fitopatol Bras 30:S 185 Ramalho TO, Figueira AR, Sotero AJ, Wang R, Geraldino Duarte PS, Farman M, Goodin MM (2014) Characterization of Coffee ringspot virus-Lavras: a model foran emerging threat to coffee production and quality. Virology 464–465:385–396 Ramalho TO, Figueira AR, Wang R, Jones O, Harris LE, Goodin MM (2016) Detection and survey of coffee ringspot virus in Brazil. Arch Virol 161:335–343 Reyes TT (1961) Seed transmission of coffee ring spot by excelsa coffee (Coffea excelsa). Plant Dis Reptr 45:185 Rodrigues JCV, Rodriguez CM, Moreira L, Vilalobos W, Rivera C, Childers CC (2002) Occurrence of Coffee ring spot virus, Brevipalpus mite-borne virus in coffe in Costa Rica, Central America. Plant Dis 86:564

Coffea ebracteolata Synonyms Psilanthus ebracteolatus Family: Rubiaceae

Commercial crop

Coffee ringspot dichorhavirus Taxonomic position Genus: Dichorhavirus

(CoRSV)

Family: Rhabdoviridae

CoRSV infection in plants of Psilanthus ebracteolatus was reported from Brazil Kitajima et al. (2011). The virus-infected plants exhibit leaves with ringspot lesions. The virus is transmitted by the mite vector, Brevipalpus phoenicis, in a persistent propagative manner, and also by mechanical sapinoculation. For more details of CoRSV, refer to Coffea Arabica.

Reference Kitajima EW, Chagas CM, Braghini MT, Fazuoli LC, Locali-Fabris EC, Salaroli RB (2011) Natural infection of several Coffea species and hybrids and Psilanthus ebracteolatus by the Coffee ringspot virus (CoRSV). Sci Agric (Piracicaba, Braz) 68:503–507

Coix chinensis (Coix seed) Family: Poaceae

Medicinal

Maize chlorotic mottle virus Taxonomic position Genus: Machlomovirus

(MCMV)

Family: Tombusviridae

C

614

Cola chlamydantha

MCMV infection in plants of Coix chinensis was reported from China (Huang et al. 2016). The virusinfected coix seed plants exhibit chlorotic and mosaic symptoms. The virus is transmitted by beetle vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of MCMV, refer to Zea mays.

Reference Huang MJ, Wen G, Li M, Sun C, Sun Y, Zhao M, He Y (2016) First report of Maize chlorotic mottle virus naturally infecting Sorghum and Coix seed in China. Plant Dis 100:1955

Cola chlamydantha Family: Malvaceae

Trees/shrubs

Cacao swollen shoot Togo B virus

(CSSTBV)

Synonyms Cacao swollen shoot virus (CSSV) Taxonomic position Genus: Badnavirus

Family: Caulimoviridae

CSSTBV infection in plants of Cola chlamydantha was reported from Ghana (Legg and Bonney 1967). The virus-infected Cola chlamydantha plants exhibit transient leaf chlorosis symptoms. The virus is transmitted by mealybug vectors in a semi-persistent manner. For more details of CSSTBV, refer to Theobroma cacao.

Reference Legg JT, Bonney JK (1967) The host range and vector species of viruses from Cola chlamydantha K. Schum., Adansonia digitata L. and Theobroma cacao L. Ann Appl Biol 60:399–403

Colchicum autumnale (Autumn crocus, Meadow saffron) Family: Colchicaceae

Medicinal

Meadow saffron breaking virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

(MSBV)

Coleus spp. (Coleus blumei; C. scutellarioides)

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Geographical distribution MSBV was reported on Colchicum autumnale plants from France (Poutaraud et al. 2004). Symptoms and host(s) The virus-infected autumn crocus plants showed wilting, necrosis and decay, and an obvious flower break symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, measuring 680–900 nm long and 11–13 nm wide. The genome is a single molecule of linear, positive-sense single-stranded RNA of c.9.5–10 kb. A partial genome sequence of 1625 nt is available (AY388995) (Poutaraud et al. 2004; Revers and Garcia 2015; Wylie et al. 2017).

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV was reported from Colchicum autumnale in the USA (Waterworth and Povish 1972). The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

References Poutaraud A, Desbiez C, Lemaire O, Lecoq H, Herrbach E (2004) Characterisation of a new Potyvirus species infecting meadow saffron (Colchicum autumnale). Arch Virol 149:1267–1277 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Waterworth HE, Povish WR (1972) Tobacco ringspot virus from naturally infected dogwood, autumn crocus and forsythia. Plant Dis Reptr 56:336–337 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Coleus spp. (Coleus blumei; C. scutellarioides) Family: Lamiaceae

Coleus blumei viroid 1 Taxonomic position Genus: Coleviroid

Ornamental

(CbVd-1)

Family: Pospiviroidae

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616

Coleus spp. (Coleus blumei; C. scutellarioides)

Geographical distribution CbVd-1 was first detected in commercially field-grown yellow Coleus (Coleus blumei) from Brazil (Fonseca et al. 1989), in Coleus scutellarioides from Canada (Singh et al. 1991), and in Coleus blumei from Japan (Ishiguro et al. 1996) and was subsequently found widely in the world (Li et al. 2006; Chung and Choi 2008; Adkar-Purushothama et al. 2013; Jiang et al. 2014; Roslan et al. 2017). Symptoms and host(s) The viroid-infected coleus plants exhibit discoloration and growth retardation symptoms. However, some of the cultivars are asymptomatic (Chung and Choi 2008). Transmission The viroid is transmitted from Coleus to Coleus in a mechanical manner (Fonseca et al. 1989; Singh et al. 1991; Jiang et al. 2014) and by vegetative propagation. Seed transmission is unusually high among the viroid species, and the rate is variable, from very low to even 100%, depending on the cultivars (Singh et al. 1991; Ishiguro et al. 1996; Chung and Choi 2008). CbVd-1 was transmissible experimentally to Ocimum sanctum, Mentha spicata, Mentha arvensis, Ocimum basilicum, and Melissa officinalis without developing symptoms (Ishiguro et al. 1996). Etiology and genome properties The genome consists of a single-stranded circular RNA of 248 nt, which is noncoding (X52960 = NC_003681) (Spieker et al. 1990; Spieker 1996a, b; Gora-Sochacka 2004; Flores et al. 2011; Giguere et al. 2014; Jiang et al. 2014).

Coleus blumei viroid 2 Taxonomic position Genus: Coleviroid

(CbVd-2)

Family: Pospiviroidae

Geographical distribution CbVd-2 was reported in Coleus blumei from Germany and China (Spieker 1996a; Fu et al. 2011; Jiang et al. 2014). Symptoms and host(s) Coleus is the only known host for CbVd-2, in which CbVd-2 can replicate without showing symptoms. Transmission The viroid is transmissible mechanically to Coleus (Jiang et al. 2014) and is spread in part by vegetative propagation of the host. Etiology and genome properties The genome consists of a single-stranded circular RNA of 301 nt which is noncoding (X95365 = NC_003682) (Gora-Sochacka 2004; Giguere et al. 2014; Jiang et al. 2014).

Coleus spp. (Coleus blumei; C. scutellarioides)

Coleus blumei viroid 3 Taxonomic position Genus: Coleviroid

617

(CbVd-3)

Family: Pospiviroidae

Geographical distribution CbVd-3 was reported in Coleus blumei from Germany and China (Spieker 1991, 1996a; Spieker et al. 1996; Jiang et al. 2011, 2014). Symptoms and host(s) Coleus blumei is the only known host for CbVd-3, in which CbVd-3 can replicate without showing symptoms. Transmission The viroid is transmissible mechanically to Coleus (Jiang et al. 2014) and is spread in part by vegetative propagation of the host. Etiology and genome properties The genome consists of a single-stranded circular RNA of 364 nt, which is noncoding (X95290 = NC_003683) (Spieker et al. 1996; Gora-Sochacka 2004; Giguere et al. 2014).

Coleus blumei viroid 5

(CbVd-5)

Taxonomic position CbVd-5 is a tentative member of the genus Coleviroid and family Pospiviroidae Geographical distribution CbVd-5 was reported in Coleus blumei from China, India, Malaysia, and Indonesia (Hou et al. 2009a; Jiang et al. 2013, 2014; Tsushima and Sano 2015; Roslan et al. 2017). Symptoms and host(s) The viroid was detected from asymptomatic Coleus. Transmission The viroid is transmissible mechanically by slash inoculation to Coleus (Hou et al. 2009a). This viroid is transmissible through pollen and true seed (Jiang et al. 2014) and is spread in part by vegetative propagation of the host. Etiology and genome properties The viroid consists of 274 nt that contain the central conserved region of Coleviroids (NC_012127) (Hou et al. 2009a).

Coleus blumei viroid 6

(CbVd-6)

Taxonomic position CbVd-6 is a tentative member of the genus Coleviroid and family Pospiviroidae.

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Coleus spp. (Coleus blumei; C. scutellarioides)

Geographical distribution CbVd-6 was detected in Coleus blumei from China and Japan (Hou et al. 2009b; Jiang et al. 2014). Symptoms and host(s) The viroid was detected from asymptomatic Coleus. Transmission The viroid is transmissible mechanically to Coleus (Hou et al. 2009b; Jiang et al. 2014). This viroid is transmissible through true seed but primarily through use of infected planting material (Jiang et al. 2014). Etiology and genome properties The viroid consists of 342 nt that containsthe central conserved region of Coleviroids (NC_012805) (Hou et al. 2009b).

Coleus vein necrosis virus Taxonomic position Genus: Carlavirus

(CVNV)

Family: Betaflexiviridae

Geographical distribution CVNV infection in plants of Coleus spp. was reported from the USA and Israel (Mollov et al. 2007). Symptoms and host(s) This virus produces veinal necrosis in coleus plants. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is readily transmissible by mechanical sap-inoculation to coleus and Nicotiana spp. Virion properties and genome The virions are flexuous filaments about 640 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8727 nt (NC_009764) and includes six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Adams et al. 2004; Mollov et al. 2007).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Coleus spp. was reported from Louisiana, USA (Holcomb and Valverde 1991; Shahmohammadi et al. 2015). The virus-infected plants exhibit symptoms of mosaic, oak-leaf line patterns, ringspots, and color fading. The virus is transmitted by a number of aphid species in a non-

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persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

C Family: Tospoviridae

INSV was reported to infect Coleus spp. plants in the USA (Dobhal et al. 2014) and Europe (Roggero et al. 1999). The virus-infected coleus plants exhibit symptoms of ringspots, arc-shaped lesions, and irregular line patterns which are the easiest to observe on cultivars with light-colored leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sapinoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tobacco etch virus Taxonomic position Genus: Potyvirus

(TEV)

Family: Potyviridae

TEV infection in plants of Coleus blumei was reported from the USA (Lockhart et al. 2010). The virusinfected coleus plants exhibit necrotic foliar symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TEV, refer to Nicotiana tabacum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV was reported to infect Coleus spp. from Greece and the USA (Hausbeck et al. 1992; Chatzivassiliou et al. 2000; Parrella et al. 2003; Dobhal et al. 2014). The virus-infected coleus plants show ringspots, arc-shaped lesions, and irregular line patterns. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Adkar-Purushothama CR, Nagaraja H, Sreenivasa MY, Sano T (2013) First report of Coleus blumei viroid infecting coleus in India. Plant Dis 97:149 Chatzivassiliou EK, Livieratos I, Jenser G, Katis NI (2000) Ornamental plants and thrips populations associated with Tomato spotted wilt virus in Greece. Phytoparasitica 28:257–264

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Chung BN, Choi GS (2008) Incidence of Coleus blumei viroid 1 in seeds of commercial coleus in Korea. Plant Pathol J 24:305–308 Dobhal S, Arif M, Olson J, Mendoza-Yerbafría A, Aguilar-Moreno S, Perez-Garcia M, Ochoa-Corona FM (2014) Sensitive detection and discrimination method for studying multiple infections of five major plant viruses infecting ornamental plants in nursery environments. Ann Appl Biol 166:286–296 Flores R, Daros J, Hernandez J-A, Di Serio F (2011) Viroids. In: Encyclopedia of life sciences (ELS). John Wiley and Sons, Ltd, Chichester. https://doi.org/10.1002/9780470015902.a0000434.pub3 Fonseca MEN, Boiteux LS, Singh RP, Kitajima EW (1989) A small viroid in coleus species from Brazil. Fitopatol Bras 14:94–96 Fu FH, Li SF, Jiang DM, Wang HQ, Liu AQ, Sang LW (2011) First report of Coleusblumei viroid 2 from commercial coleus in China. Plant Dis 95:494 Giguere T, Raj Adkar-Purushothama C, Perreault J-P (2014) Comprehensive secondary structure elucidation of four genera of the family Pospiviroidae. PLoS One 9(6):e98655. https://doi.org/10.1371/journal.pone.0098655 Gora-Sochacka A (2004) Viroids: unusual small pathogenic RNAs. Acta Biochim Pol 51(3):587–607 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among greenhouse ornamentals in Pennsylvania. Plant Dis 76:795–800 Holcomb GE, Valverde RA (1991) Identification of a virus causing a mosaic on coleus. Plant Dis 75:1183–1185 Hou WY, Sano T, Li SF, Li F, Li L, Wu ZJ (2009a) Identification and characterization of a new coleviroid (CbVd-5). Arch Virol 154:315–320 Hou WY, Li SF, Wu ZJ, Jiang DM, Sano T (2009b) Coleus blumei viroid 6: anew tentative member of the genus Coleviroid derived from natural genome shuffling. Arch Virol 154:993–997 Ishiguro A, Sano T, Harada Y (1996) Nucleotide sequence and host range of coleus viroid isolated from coleus (Coleus blumei Benth.) in Japan. Ann Phytopathol Soc Japan 62:84–86 Jiang DM, Li SF, Fu FH, Wu ZJ, Xie LH (2011) First reported occurrence of Coleus blumei viroid 3 from Coleus blumei in China. J Plant Pathol 93:S4.63–S4.89 Jiang DM, Li SF, Fu FH, Wu ZJ, Xie LH (2013) First report of Coleus blumei viroid 5 from Coleus blumei in India and Indonesia. Plant Dis 97:561 Jiang D, Gao R, Qin L, Wu Z, Xie L, Hou W, Li S (2014) Infectious cDNA clones of four viroids in Coleus blumei and molecular characterization of their progeny. Virus Res 180:97–101 Li S-F, Su Q, Guo R, Tsuji M, Sano T (2006) First report of Coleus blumei viroid from coleus in China. Plant Pathol 55:565 Lockhart BEL, Mason SL, Johnson DA, Mollov DS (2010) First report of Tobacco etch virus infection in coleus in the United States. Plant Dis 94:921 Mollov DN, Eichstaedt KA, Beckman NG, Daughtrey ML, Lockhart BE (2007) Identification and characterization of a Carlavirus causing veinal necrosis of coleus. Plant Dis 91:754–757 Parrella G, Gognalons P, Gebre-Selassi K, Vovlas C, Marchoux G (2003) An update of the host range of Tomato spotted wilt virus. J Plant Pathol 85(4):227–264 Roggero P, Ciuffo M, Dellavalle G, Gotta P, Gallo S, Peters D (1999) Additional ornamental species as hosts of Impatiens necrotic spot tospovirus in Italy. Plant Dis 83:967 Roslan NNC, Thanarajoo SS, Kadir J, Kong LL, Vadamalai G (2017) First report of Coleus blumei viroid in Malaysia. J Plant Pathol 99:800 Shahmohammadi N, Dizadji A, Habibi MK, Nateqi M (2015) First report of Cucumber mosaic virus infecting Bougainvillea spectabilis, Coleus blumei, Kalanchoe blossfeldiana and Zinnia elegans in Iran. J Plant Pathol 97(2):394 Singh RP, Boucher A, Singh A (1991) High incidence of transmission and occurrence of a viroid in commercial seeds of Coleus in Canada. Plant Dis 75:184–187 Spieker RL (1991) A new class of viroids in Coleus blumei: sequencing, cloning structure/function-analysis by sitedirected mutagenesis and expression in Nicotiana tabacum. Karlsruhe Contrib Plant Physiol 22:1–226 Spieker RL (1996a) In vitro-generated “inverse” chimeric Coleus blumei viroids evolve in vivo into infectious RNA replicons. J Gen Virol 77:2839–2846 Spieker R (1996b) A new sequence variant of Coleus blumei viroid 1 from the Coleus blumei cultivar “Rainbow Gold”. Arch Virol 141:2153–2161 Spieker RL, Haas B, Charng YC, Freimüller K, Sänger HL (1990) Primary and secondary structure of a new viroid ‘species’ (CbVd1) present in the Coleus blumei cultivar Bienvenue’. Nucleic Acids Res 18:3998 Spieker RL, Marinkovic S, Sänger HL (1996) A new sequence variant of Coleus blumei viroid 3 from the Coleus blumei cultivar ‘Fairway Ruby’. Arch Virol 141:1377–1386 Tsushima T, Sano T (2015) First report of Coleus blumei viroid 5 infection in vegetatively propagated clonal coleus cv. ‘Aurora black cherry’ in Japan. New Dis Rep 32:7

Colocasia esculenta (Dasheen or Taro or Cocoyam)

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Colocasia esculenta (Dasheen or Taro or Cocoyam) Family: Araceae

Tuber crop

Colocasia bobone disease-associated cytorhabdovirus Taxonomic position Genus: Cytorhabdovirus

(CBDaV)

Family: Rhabdoviridae

Geographical distribution CBDaV infection in plants of Colocasia esculenta was reported from Solomon Islands and Papua New Guinea (Higgins et al. 2016). Symptoms and host(s) The virus-infected taro plants exhibit symptoms of severe stunting and gall formation and associated with bobone disease. Transmission Colocasia bobone disease virus (CBDV) is known to be transmitted by the planthopper Tarophagus proserpina (James et al. 1973), but it is not clear that CBDaV is the same as CBDV, and the full genome sequence shows that CBDaV has only a single ORF between the P and M coding sequences, similar to the situation in aphid-transmitted cytorhabdoviruses, whereas other known planthopper-transmitted cytorhabdoviruses have multiple ORFs between their P and M coding sequences (Higgins et al. 2016). This suggests that CBDaV may be transmitted by aphids. Virion properties and genome The virions are enveloped and bacilliform and measure 73 nm in diameter and 269 nm long. There are distinct surface projections (spikes) dispersed evenly over all the surface. The genome is a monopartite, negative-sense, single-stranded RNA of 12,193 nt is unsegmented (KT381973 = NC_034551) and has six ORFs in the anti-genomic strand equivalent to putative (3–50 ) N, P, P3, M, G, and L genes (Pearson et al. 1999; Dietzgen 2011; Higgins et al. 2016; Walker et al. 2018).

Colocasia bobone disease virus

(CBDV)

Taxonomic position CBDV is a tentative member of the family Rhabdoviridae. Geographical distribution CBDV infection in plants of Colocasia esculenta was reported from Papua New Guinea, Solomon Islands, and Pacific Islands (James et al. 1973; Jackson and Gollifer 1975; Rodoni et al. 1994; Pearson et al. 1999; Revill et al. 2005b). The disease in taro caused by CBDV alone is called bobone.

C

622

Colocasia esculenta (Dasheen or Taro or Cocoyam)

Symptoms and host(s) In taro this virus is characterized by the development of mosaic symptoms, severe stunting, and distorted, thickened, brittle leaves. Galls may be present on the petioles and sometimes on the larger veins. The virus also exhibits leaf twisting and curling and plants are stunted. Plants sometimes recover and grow normally (Gollifer et al. 1977). Transmission The virus is transmitted by the taro planthopper vector, Tarophagus proserpina, in a persistent manner (James et al. 1973; Shaw et al. 1979). The virus is not transmissible by mechanical sap-inoculation, and the host range is limited to aroids. The virus is not transmissible by contact between plants, not transmitted by seed, and not transmitted by pollen. The use of virus-infected vegetative planting material (corms) is the main mode of virus spread. Virion properties and genome The virions are rhabdo- or bullet-shaped, with a clear modal length of 300–335 nm and 50–55 nm wide (James et al. 1973). The genome is a linear unipartite negative-sense ssRNA.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Colocasia esculenta was reported from China (Wang et al. 2014). The virusinfected taro plants exhibit symptoms of leaf chlorosis or chlorotic spots and feather-like mosaic symptom on their leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Dasheen mosaic virus Taxonomic position Genus: Potyvirus

(DsMV)

Family: Potyviridae

Geographical distribution DsMV infection in plants of Colocasia esculenta was first reported from the USA (Zettler et al. 1970). The virus occurs worldwide (Alconero and Zettler 1971; Abo El-Nil and Zettler 1976; Zettler and Hartman 1987; Zettler et al. 1987; Hu et al. 1994, 1995; Ram et al. 2003; Kim et al. 2004; Revill et al. 2005b; Babu et al. 2011; Babu and Hegde 2014; Wang et al. 2017a). Symptoms and host(s) DsMVon taro plants produces conspicuous foliar mosaic, mottle, and chlorotic “feathering” symptoms that are apparent. An individual virus-infected plant may have leaves with pale green feathering symptoms, whereas others may have severe leaf curling and malformation or slight vein-banding symptoms or no visible symptoms at all. Foliar symptoms are intermittently expressed. Severity and persistence of expressed symptoms vary according to plant genotype. Growth of virus-infected plants is retarded, and corms do not exhibit symptoms. In DsMV-infected taro, two or three leaves may show symptoms, and then apparently healthy leaves are produced, or leaves may alternate between

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asymptomatic and symptomatic on the same plant (Elliott et al. 1997). Plants sometimes recover from the symptoms and produce leaves healthy in appearance. This virus infects crops such as Aglaonema, Caladium, Dieffenbachia, Xanthosoma, and Zantedeschia spp. (Chen and Li 1996). Transmission The virus is transmitted by aphid vectors, Myzus persicae, Aphis craccivora, and Aphis gossypii, in a non-persistent manner. The aphid, Pentalonia nigronervosa, however, is apparently not a vector (Brunt et al. 1990; Gollifer et al. 1977). The virus is transmissible by mechanical sap-inoculation, and has a narrow experimental host range outside Araceae, infecting plants such as Tetragonia expansa, Chenopodium spp., Nicotiana spp., and Saponaria vaccaria (Babu et al. 2011). The virus is vegetatively transmissible by suckers, corms, and infected cuttings. The virus is not transmissible by contact between plants. The virus is not seed and pollen transmissible (Nelson 2008). Virion properties and genome The virions are non-enveloped, flexuous filaments measuring 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 10,038 nt (AJ298033 = NC_003537) (Revers and Garcia 2015). The genome contains single large open reading frame (ORF) with AUG initiation codon at 168–170 nts and UAA termination codon at 9741–9743 nts followed by 30 - UTR of 248 nt. The 50 - UTR is 167 nt. The deduced poly protein of this ORF contains 3162 amino acids with a calculated molecular weight of 362.5 kDa (Brunt et al. 1990; Kim et al. 2004; Revers and Garcia 2015; Wylie et al. 2017).

Groundnut bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(GBNV)

Family: Tospoviridae

GBNV infection in plants of Colocasia esculenta was reported from India (Sivaprasad et al. 2011). The virus-infected taro plants exhibit symptoms of mosaic, chlorotic spotting, and necrotic flecking of leaves. The virus is transmitted by a thrips vector, Thrips palmi, in a persistent-propagative manner, and also by mechanical sap-inoculation to a wide range of host plants. The use of infected vegetative planting material is responsible for the primary spread of this virus disease. For more details of GBNV, refer to Arachis hypogaea.

Konjac mosaic virus Taxonomic position Genus: Potyvirus

(KoMV)

Family: Potyviridae

KoMV infection in plants of Colocasia esculenta plants was reported from Japan, Taiwan, and India (Padmavathi et al. 2011, 2015). The virus-infected taro plants exhibit symptoms of mosaic, chlorotic feathery mottling, chlorotic spots, leaf deformation, and chlorotic ringspots. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of KoMV, refer to Amorphophallus paeoniifolius.

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624

Colocasia esculenta (Dasheen or Taro or Cocoyam)

Taro bacilliform CH virus Taxonomic position Genus: Badnavirus

(TaBCHV)

Family: Caulimoviridae

Geographical distribution TaBCHV infection in plants of Colocasia esculenta was reported from China, the USA and the East African countries of Ethiopia, Kenya, Tanzania and Uganda (Kazmi et al. 2015; Wang et al. 2017b, c; Kidanemariam et al. 2018). Symptoms and host(s) The virus-infected taro plants show mild feathery mosaic symptoms on young leaves and brown spots on matured leaves. Transmission The virus is transmissible by the use of infected planting material. No vector has been identified to date. Virion properties and genome The virions are bacilliform in shape with a diameter of 30 nm and modal particle length is 130 nm. The genome is a monopartite, circular double-stranded DNA of 7641 bp (KP710178) with a single-stranded discontinuity at one site in each strand. The genome contains six open reading frames (ORFs) (Olszewski and Lockhart 2011; Kazmi et al. 2015; Bhat et al. 2016; Wang et al. 2017c).

Taro bacilliform virus Taxonomic position Genus: Badnavirus

(TaBV)

Family: Caulimoviridae

Geographical distribution TaBV was first observed on Colocasia esculenta from Solomon Islands (James et al. 1973). The virus spreads in Asia, French Polynesia, New Caledonia, Papua New Guinea, Samoa, Solomon Islands, Cook Islands, Fiji, Vanuatu and the East African countries of Kenya, Tanzania and Uganda (Jackson and Gollifer 1975; Revill et al. 2005b; Macanawai 2003; Harding 2008; Aarthy et al. 2017; Kidanemariam et al. 2018). Symptoms and host(s) The virus-infected taro plants generally remain symptomless. However where symptoms do occur, they are usually mild and include stunting, mild mosaic, and downcurling of the leaves. Some transient veinclearing and slight leaf malformation symptoms are also observed. Rarely, vein chlorosis occurs throughout the leaf (Yang et al. 2003a). In addition to C. esculenta, the natural host range of this virus includes Alocasia macrorrhiza and Xanthosoma spp. Transmission The virus is transmitted by mealybug vectors, Planococcus citri, Pseudococcus solomonensis, and Pseudococcus longispinus in a semi-persistent manner (Gollifer et al. 1977) Transmission of virus has been reported even through true seeds and pollen of taro (Macanawai et al. 2005). The virus is not

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mechanically sap-transmissible and not transmitted by contact between plants. The use of virusinfected vegetative planting material (corms) is the main mode of the virus spread. Virion properties and genome The virions are bacilliform with a length of 130 nm and a width of 30 nm (James et al. 1973). The genome is a monopartite circular double-stranded DNA of 7458 bp (NC_004450). The genome contains four open reading frames (ORFs) on the plus strand that potentially encode proteins of 17, 16, 214, and 13 kDa (Yang et al. 2003b; Olszewski and Lockhart 2011; Bhat et al. 2016).

Taro vein chlorosis nucleorhabdovirus Taxonomic position Genus: Nucleorhabdovirus

(TaVCV)

Family: Rhabdoviridae

Geographical distribution TaVCV infection in plants of Colocasia esculenta was found in several Pacific countries including Fiji, Republic of Palau, Solomon Islands, Tuvalu, Vanuatu, New Caledonia, Federated States of Micronesia, the USA, and New Zealand (James et al. 1973; Pearson et al. 1999; Revill et al. 2005a, b; Long et al. 2014). Symptoms and host(s) The virus-infected taro plants exhibit symptoms of distinct vein chlorosis particularly at the leaf margins which spreads between the veins as the leaves mature and leads to necrosis of affected tissues. No galls are present on the leaf blades and petioles, and plants are not stunted. Transmission The virus is not mechanically sap-transmissible. The use of plantings from infected plants is the main source of the spread. No vector has been identified. Virion properties and genome The virions are enveloped and bacilliform and measure 70 nm in diameter and 210 nm long (Pearson et al. 1999). There are distinct surface projections (spikes) dispersed evenly over all the surface. The genome is a monopartite, negative-sense, single-stranded RNA of 12,020 nt is unsegmented (AY674964 = NC_006942). The genome contains six open reading frames (ORFs) in the antigenomic sequence, equivalent to nucleocapsid (N) gene, phosphoprotein (P) gene, gene 3, matrix protein (M) gene, glycoprotein (G) gene, and polymerase protein (L) gene which have been identified (Revill et al. 2005a; Jackson et al. 2011; Walker et al. 2018).

References Aarthy MB, Arutselvan R, Devi A, Jeeva ML, Makesh Kumar T (2017) Molecular characterization of Taro bacilliform virus infecting taro (Colocasia esculena) in India. In: 26th Annual Conference of Indian Virological Society (VIROCON 2017), Mangaluru, India. p 95 Abo El-Nil MM, Zettler FW (1976) Natural occurrence of Dasheen mosaic virus in Egyptian taro, Colocasia esculenta. Plant Dis Reptr 60:281–285

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Colocasia esculenta (Dasheen or Taro or Cocoyam)

Alconero R, Zettler FW (1971) Virus infections of Colocasia and Xanthosoma in Puerto Rico. Plant Dis Reptr 55:506–508 Babu B, Hegde V (2014) Molecular characterization of Dasheen mosaic virus isolates infecting edible aroids in India. Acta Virol 58(1):34–42 Babu B, Hegde V, Makeshkumar T, Jeeva ML (2011) Detection and identification of Dasheen mosaic virus infecting Colocasia esculenta in India. Indian J Virol 22(1):59–62 Bhat AI, Hohn T, Selvarajan R (2016) Badnaviruses: the current global scenario. Viruses 8(6):177 Brunt A, Crabtree K, Gibbs A (1990) Viruses of tropical plants. CAB International and ACIAR, Wallingford 707 pp Chen J, Li D (1996) Dasheen mosaic virus from thirteen Araceae crops. Acta Microbiol Sin 36:126–131 Dietzgen RG (2011) Cytorhabdovirus. Rhabdoviridae. In: The Springer index of Viruses. Springer, New York, pp 1709–1713. https://doi.org/10.1007/978-0-387-95919-1_277 Elliott MS, Zettler FW, Brown LG (1997) Dasheen mosaic potyvirus of edible and ornamental aroids, Plant Pathology Circular No. 384. Fla Dept and Consumer Services., July/August 1997, Florida Gollifer DE, Jackson GVH, Dabek AJ, Plumb RT, May YY (1977) The occurrence and transmission of viruses of edible aroids in the Solomon Islands and the Southwest Pacific. Pestic Artic News Summ 23:171–177 Harding R (2008) Taro bacilliform virus. In: Rao GP, Paul Khurana SM, Lenardon SL (eds) Characterization, diagnosis and management of plant viruses, Industrial Crops, vol I. Studium Press LLC, Texas, pp 341–348 Higgins CM, Bejerman N, Li M, James AP, Dietzgen RG, Pearson MN, Revill PA, Harding RM (2016) Complete genome sequence of Colocasia bobone disease-associated virus, a putative Cytorhabdovirus infecting taro. Arch Virol 161:745–748 Hu JS, Wang M, Reolanei R, Meleisea S (1994) Detection of Dasheen mosaic virus from taro plants in the field and in tissue culture. Plant Dis 78:754 Hu JS, Meleisea S, Wang M, Shaarawy MA, Zettler FW (1995) Dasheen mosaic potyvirus in Hawaiian taro. Australas Plant Pathol 24:112–117 Jackson GVH, Gollifer DE (1975) Disease and pest problems of taro (Colocasia esculenta L. Schott) in the British Islands. Pesticide articles and news. Summaries 21:45–53 Jackson AO, Goodin MM, Ganesan U, Bragg J (2011) Nucleorhabdovirus. Rhabdoviridae. In: The Springer index of Viruses. Springer, New York, pp 1741–1745. https://doi.org/10.1007/978-0-387-95919-1_281 James M, Kenten RH, Woods RD (1973) Virus-like particles associated with two diseases of Colocasia esculenta (L.) Schott in the British Solomon Islands. J Gen Virol 21:145–153 Kazmi SA, Yang Z, Hong N, Wang G, Wang Y (2015) Characterization by small RNA sequencing of Taro bacilliform CH virus (TaBCHV), a novel Badnavirus. PLoS One 10:E0134147 Kidanemariam DB, Sukal AC, Abraham AD, Stomeo F, Dale JL, James AP, Harding RM (2018) Identification and molecular characterization of Taro bacilliform virus and Taro bacilliform CH virus from East Africa. Plant Pathol. https://doi.org/10.1111/ppa.12921 Kim MK, Kwon SB, Yoon JY, Ryu KH, Heo SJ, Hong JK, Kim KH, Choi JK (2004) Characterization of a Korean isolate of Dasheen mosaic virus isolated from taro (Colocasia esculenta Schott) in Korea. Plant Pathol J 20:135–141 Long MH, Ayin C, Li R, Hu JS, Melzer MJ (2014) First report of Taro vein chlorosis virus infecting taro (Colocasia esculenta) in the United States. Plant Dis 98:1160–1161 Macanawai A (2003) The epidemiology of Taro bacilliform virus in Samoa. MSc Thesis. The University of the South Pacific, Samoa Macanawai AR, Ebenebe AA, Hunter D, Devitt LC, Hafner GJ, Harding RM (2005) Investigations into the seed and mealybug transmission of Taro bacilliform virus. Aust Plant Pathol 34:73–76 Nelson SC (2008) Dasheen mosaic of edible and ornamental aroids, Plant Disease CTAHR. University of Hawaii, Honolulu Available at: http://www.ctahr.hawaii.edu/oc/freepubs/pdf/PD-44.pdf Olszewski NE, Lockhart B (2011) Badnavirus. Caulimoviridae. In: The Springer index of Viruses. Springer, New York, pp 263–269. https://doi.org/10.1007/978-0-387-95919-1_40 Padmavathi M, Srinivas KP, Subba Reddy CV, Ramesh B, Navodayam K, Krishnaprasadji J, Babu Ratan P, Sreenivasulu P (2011) Konjac mosaic virus naturally infecting three aroid plant species in Andhra Pradesh, India. J Phytopathol 159:133–135 Padmavathi M, Srinivas KP, Hema M, Sreenivasulu P (2015) Development of duplex RT-PCR for detection of Konjac mosaic virus and Spathyphyllum chlorotic vein banding virus in taro and peace lily. Indian J Biotechnol 14:120–122 Pearson MN, Jackson GVH, Saelea J, Morar SG (1999) Evidence for two rhabdoviruses in taro (Colocasia esculenta) in the Pacific region. Australas Plant Pathol 28:248–253 Ram R, Joshi A, Verma N, Kulshrestha S, Raikhy G, Hallan V, Zaidi AA (2003) First report of Dasheen mosaic virus infecting four ornamental aroids in India. Plant Pathol 52:411 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Revill P, Trinh X, Dal J, Harding R (2005a) Taro vein chlorosis virus: characterization and variability of a new Nucleorhabdovirus. J Gen Virol 86:491–499

Columnea spp.

627

Revill PA, Jackson GVH, Hafner GJ, Yang I, Maino MK, Dowling ML, Devitt LC, Dale JL, Harding RM (2005b) Incidence and distribution of viruses of taro (Colocasia esculenta) in Pacific Island countries. Aust Plant Pathol 34:327–331 Rodoni BC, Dale JL, Harding RM (1994) Review of Alomae disease of taro. P N G J Agric For Fish 37:14–18 Shaw DE, Plumb RT, Jackson GVH (1979) Virus diseases of taro (Colocasia esculenta) and Xanthosoma spp. in Papua New Guinea. P N G Agric J 30:71–97 Sivaprasad Y, Bhakara Reddy BV, Naresh Kumar CVM, Raja Reddy K, Sai Gopal DVR (2011) First report of Groundnut bud necrosis virus infecting taro (Colocasia esculenta). Aust Plant Dis Notes 6:30–32 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Rhabdoviridae. J Gen Virol 99:447–448 Wang YF, Wang GP, Wang LP, Hong N (2014) First report of Cucumber mosaic virus in taro plants in China. Plant Dis 98(4):574–575 Wang Y, Wu B, Borth WB, Hamim I, Green JC, Melzer MJ, Hu JS (2017a) Molecular characterization and distribution of two strains of Dasheen mosaic virus on taro in Hawaii. Plant Dis 101:1980–1989 Wang YN, Hu JS, Borth WB, Hamim I, Green JC, Melzer MJ (2017b) First report of Taro bacilliform CH virus (TaBCHV) on taro (Colocasia esculenta) in Hawaii, USA. Plant Dis 101:1334 Wang Y, Borth WB, Green JC, Hamim I, Cao K, Hu JS, Melzer MJ (2017c) Genome characterization and distribution of Taro bacilliform CH virus on taro in Hawaii, USA. Eur J Plant Pathol 150:1107–1111 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Yang IC, Hafner GJ, Revill TA, Dale JL, Harding RM (2003a) Sequence diversity of south pacific isolates of Taro bacilliform virus and development of PCR based diagnositc test. Arch Virol 148:1957–1968 Yang IC, Hafner GJ, Revill PA, Dale JL, Harding RM (2003b) Genomic characterisation of Taro bacilliform virus. Arch Virol 148:937–949 Zettler FW, Hartman RD (1987) Dasheen mosaic virus as pathogen of cultivated aroids and control of the virus by tissue culture. Plant Dis 71:958–963 Zettler FW, Foxe MJ, Hartman RD, Edwardson JR, Christie RG (1970) Filamentous viruses infecting dasheen and other araceous plants. Phytopathology 60:983–987 Zettler FW, Tsai JH, Faan HC, Ke C, Lu KC (1987) Dasheen mosaic virus infecting taro in People’s Republic of China. Plant Dis 71:837–839

Columnea spp. Family: Gesneriaceae

Columnea latent viroid Taxonomic position Genus: Pospiviroid

Ornamental

(CLVd)

Family: Pospiviroidae

Geographical distribution CLVd was detected in plants of Columnea erythrophae from the USA (Owens et al. 1978; Hammond et al. 1989). Symptoms and host(s) The viroid-infected Columnea plants do not exhibit any symptoms. The viroid was first detected from Columnea and then found in various ornamental plants, such as Brunfelsia, Nematanthus wettsteinii, and Gloxinia (Matsushita and Tsuda 2015).

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Columnea spp.

Transmission The viroid is transmissible mechanically to Gynura aurantiaca, cucumber, tomato, and potato (Owens et al. 1978). Although CLVd infects Columnea asymptomatically, the infection of tomato and potato results in severe disease symptoms similar to those incited by PSTVd, suggesting that Columnea can serve as a potential reservoir for serious disease for susceptible crops. Etiology and genome properties The genome consists of a single-stranded circular RNA of 350–374 nts that contains a chimeric structure with properties of the genera Pospiviroid and Hostuviroid (X15663 = NC_003538), namely, CLVd shares the central conserved region (CCR, the main structural feature adopted for genus demarcation within the family Pospiviroidae) with HpSVd (the genus Hostuviroid); however, CLVd also has the terminal conserved region (TCR) which is present in all pospi- and apscaviroids but absent from HpSVd. From a biological point of view, CLVd shares with pospiviroids the ability to infect solanaceous hosts, inducing symptoms similar to those caused by most members of this genus. Therefore, CLVd is currently classified in the genus Pospiviroid in accordance with phylogenetic analysis (Hammond et al. 1989; Gora-Sochacka 2004; Di Serio et al. 2014; Giguere et al. 2014).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Columnea spp. was reported from the Czech Republic (Mertelik et al. 2000, 2002). The virus-infected columnea plants show chlorotic spots and pattern symptoms. The virus is transmitted by a thrips vector Frankliniella occidentalis in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV was reported to infect Columnea spp. in the USA (Zettler and Nagel 1983). The symptoms on the foliage of infected columnea plants were inconspicuous. There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Columnea spp. was reported from the Netherlands and the Czech Republic (Mertelik et al. 1996; Parrella et al. 2003). The virus-infected columnea plants exhibit distortion, mottle mosaic, necrosis, ringspots, chlorosis, and bronzing symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of

Combretum confertum (Thailand powder puff)

629

herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Di Serio F, Flores R, Verhoeven JTJ, Li S-F, Pallas V, Randles JW, Sano T, Vidalakis G, Owens RA (2014) Current status of viroid taxonomy. Arch Virol 159:3467–3478 Giguere T, Raj Adkar-Purushothama C, Perreault J-P (2014) Comprehensive secondary structure elucidation of four genera of the family Pospiviroidae. PLoS One 9(6):e98655. https://doi.org/10.1371/journal.pone.0098655 Gora-Sochacka A (2004) Viroids: unusual small pathogenic RNAs. Acta Biochim Pol 51(3):587–607 Hammond R, Smith DR, Diener TO (1989) Nucleotide sequence and proposed secondary structure of Columnea latent viroid: a natural mosaic of viroid sequences. Nucleic Acids Res 17:10083–10094 Matsushita Y, Tsuda S (2015) Host ranges of Potato spindle tuber viroid, Tomato chlorotic dwarf viroid, Tomato apical stunt viroid, and Columnea latent viroid in horticultural plants. Eur J Plant Pathol 141:193–197 Mertelík J, Götzová B, Mokrá V (1996) Epidemiological aspects of Tomato spotted wilt virus infection in the Czech Republic. Acta Hortic 432:368–375 Mertelik J, Mokra V, Gotzova B, Gabrielova S (2000) First report of Impatiens necrotic spot virus in the Czech Republic. Plant Dis 84:1045 Mertelik J, Mokra V, Gotzova B, Gabrielova S (2002) Occurrence and identification of Impatiens necrotic spot tospovirus in the Czech Republic. Acta Hortic 568:79–83 Owens RA, Smith DR, Diener TO (1978) Measurement of viroid sequence homology by hybridization with complementary DNA prepared in vitro. Virology 89:388–394 Parrella G, Gognalons P, Gebre-Selassi K, Vovlas C, Marchoux G (2003) An update of the host range of Tomato spotted wilt virus. J Plant Pathol 85(4, special issue):227–264 Zettler FW, Nagel J (1983) Infection of cultivated gesneriads by two strains of Tobacco mosaic virus. Plant Dis 67:1123–1125

Combretum confertum (Thailand powder puff) Family: Combretaceae

Weed host

African cassava mosaic virus Taxonomic position Genus: Begomovirus

(ACMV)

Family: Geminiviridae

ACMV infection in plants of Combretum confertum was reported from Nigeria (Ogbe et al. 2006; Alabi et al. 2008). The virus-infected Thailand powder puff plants exhibit bright mosaic symptoms. The virus is transmitted by a whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ACMV, refer to Manihot esculenta.

East African cassava mosaic virus Taxonomic position Genus: Begomovirus

(EACMV)

Family: Geminiviridae

EACMV infection in plants of Combretum confertum was reported from Nigeria (Ogbe et al. 2006; Alabi et al. 2008). The virus-infected Thailand powder puff plants exhibit bright mosaic symptoms. The

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Commelina spp. (Dayflower)

virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of EACMV, refer to Manihot esculenta.

References Alabi OJ, Ogbe FO, Bandyopadhyay R, Lava Kumar P, Dixon AGO, Hughes J’A, Naidu RA (2008) Alternate hosts of African cassava mosaic virus and East African cassava mosaic Cameroon virus in Nigeria. Arch Virol 153:1743–1747 Ogbe FO, Dixon AGO, Hughes JA, Alabi OJ, Okechukw R (2006) Status of cassava begomo viruses and their new natural hosts in Nigeria. Plant Dis 90:548–553

Commelina spp. (Dayflower) Family: Commelinaceae

Weed host

Brome mosaic virus

(BMV)

Taxonomic position Genus: Bromovirus

Family: Bromoviridae

BMV infection in plants of Commelina spp. was reported from Finland, South Africa, the Central USA, Canada, Russia, the former USSR, and the former Yugoslavia (Valverde 1983). The virus-infected dayflower plants exhibit mosaic symptoms. The virus is transmitted by chrysomelid beetle vectors and also by mechanical sap-inoculation. For more details of BMV, refer to Bromus spp.

Citrus leprosis virus C Taxonomic position Genus: Cilevirus

(CiLV-C)

Family: Kitaviridae

CiLV-C infection in plants of Commelina benghalensis was reported from Brazil (Nunes et al. 2012). The virus-infected dayflower plants exhibit symptoms of necrotic spots with a yellow halo in green leaves and green spots with necrotic center in senescent leaves. The virus is transmitted by the mite vector, Brevipalpus phoenicis, and also by mechanical sap-inoculation. For more details of CiLV-C, refer to Citrus spp.

Commelina mild mosaic virus

(ComMMV)

Taxonomic position ComMMV is a tentative member of the genus Potyvirus and family Potyviridae Geographical distribution ComMMV infection in plants of Commelina cyanea was reported from Australia (Zheng et al. 2011).

Commelina spp. (Dayflower)

631

Symptoms and host(s) The virus-infected plants exhibit mild mosaic leaf symptoms. Transmission Possibly the virus is transmitted by aphid vectors and also by mechanical sap-inoculation. Virion properties and genome The virus has filamentous virions. The genome consists of a single molecule of positive-sense singlestranded RNA of c.9.5–10 kb. A partial sequence of three terminal regions comprising of 1625 nt is available (HQ225836) (Zheng et al. 2011; Wylie et al. 2017).

Commelina mosaic virus Taxonomic position Genus: Potyvirus

(ComMV)

Family: Potyviridae

Geographical distribution ComMV infection was first reported in Commelina spp. from Florida, USA, by Morales and Zettler (1977). The virus spreads in the USA (Baker and Zettler 1988). Symptoms and host(s) The virus-infected dayflower plants exhibit broad, sharply defined mosaic patterns, stunting, and foliar distortion (Morales and Zettler 1977). Transmission The virus is transmitted by aphid vectors, Aphis gossypii and Myzus persicae, in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments measuring 707–808 nm long and 11–13 nm wide (Morales and Zettler 1977). The genome consists of a single molecule of linear, positive-sense ssRNA of c.9.5–10 kb (Revers and Garcia 2015; Wylie et al. 2017).

Commelina yellow mottle virus Taxonomic position Genus: Badnavirus

(ComYMV)

Family: Caulimoviridae

Geographical distribution ComYMV first reported in plants of Commelina diffusa from Guadeloupe by Migliori and Lastra (1978). The virus spreads in Guadeloupe and Sudan (Migliori and Lastra 1978; B.E.L. Lockhart unpublished data; Medberry et al. 1990; Cheng et al. 1998). Transmission The virus is likely transmissible by mealybugs; it is not transmitted by mechanical sap-inoculation. The virus is transmissible by grafting.

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Commelina spp. (Dayflower)

Virion properties and genome The virions are bacilliform and non-enveloped measuring133 nm in length and 24 nm wide. The genome consists of a circular double-stranded DNA of 7489 bp (NC_001343) with the three ORFs capable of encoding proteins of 23, 15, and 216 kD (Medberry et al. 1990; Cheng et al. 1998; Olszewski and Lockhart 2011; Bhat et al. 2016).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Commelina spp. is known worldwide (Baker and Zettler 1988; Choi et al. 1990; Duarte et al. 1994). The virus-infected dayflower plants exhibit mottling with circular areas having a sharply defined yellow border. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Commelina benghalensis was reported from Australia (Sharman et al. 2015). The virus-infected dayflower plants exhibit chlorotic mottle and tip necrosis symptoms. The virus is transmitted by the thrips vectors; the virus present in/on pollen enters into the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Yam mosaic virus Taxonomic position Genus: Potyvirus

(YMV)

Family: Potyviridae

YMV infection in plants of Commelina spp. was reported from Lome, Togo (West Africa), by Gumedzoe (1993). The virus-infected dayflower plants exhibit mosaic mottling symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of YMV, refer to Dioscorea spp.

References Baker CA, Zettler FW (1988) Viruses infecting wild and cultivated species of the Commelinaceae. Plant Dis 72:513 Bhat AI, Hohn T, Selvarajan R (2016) Badnaviruses: the current global scenario. Viruses 8(6):177 Cheng CP, Tzafrir I, Lockhart BEL, Olszewski NE (1998) Tubules containing virions are present in plant tissues infected with Commelina yellow mottle badnavirus. J Gen Virol 79(4):925–929

Conium maculatum (Hemlock or poison hemlock)

633

Choi JK, Kwon SB, Lee SY, Park WM (1990) Some properties of two isolates of Cucumber mosaic virus isolated from Aster yomena Makino and Commelina communis L. Kor J Plant Pathol 6:138–143 Duarte LML, Rivas EB, Alexandre MAV, Ferrari JT (1994) Detection of CMV isolates from Commelinaceae species. Fitopatol Bras 19:248–253 Gumedzoe MY (1993) Major virus diseases of medicinal plants in West Africa. Acta Hortic (ISHS) 331:307–310 Medberry SL, Lockhart BEL, Olszewski NE (1990) Properties of Commelina yellow mottle virus’s complete DNA sequence, genomic discontinuities and transcript suggest that it is a Pararetrovirus. Nucleic Acids Res 18:5505–5513 Migliori A, Lastra R (1978) Study of viruses on Commelina diffusa Burm. in Guadeloupe. Ann Phytopathol 10:467–477 Morales FJ, Zettler FW (1977) Characterization and electron microscopy of a Potyvirus infecting commelina. Phytopathology 67:839–843 Nunes MA, Bergamini MP, Coerini LF, Bastianel M, Novelli VM, Kitajima EW, Freitas-Astua J (2012) Citrus leprosis virus C naturally infecting Commelina benghalensis, a prevalent monocot weed of citrus orchards in Brazil. Plant Dis 96:770 Olszewski NE, Lockhart B (2011) Badnavirus. Caulimoviridae. In: The Springer index of Viruses. Springer, New York, pp 263–269. https://doi.org/10.1007/978-0-387-95919-1_40 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct Tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207 Valverde RA (1983) Brome mosaic virus isolates naturally infecting Commelina diffusa and C. communis. Plant Dis 67:1194–1196 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Zheng L, Gibbs M, Gibbs A, Rodoni B (2011) First report of a newly detected Potyvirus, Commelina mild mosaic virus, infecting Commelina spp. in Australia. Aust Plant Dis Notes 6:11–15

Conium maculatum (Hemlock or poison hemlock) Family: Apiaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Ornamental

(AMV)

Family: Bromoviridae

AMV infection in plants of Conium maculatum was reported from the USA (Howell and Mink 1981). The virus-infected poison hemlock plants exhibit calico mosaic symptoms. The virus is transmitted by a large number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Apium virus Y

(ApVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

ApVY infection in plants of Conium maculatum was reported from Australia and the USA (Moran et al. 2002; Eastwell et al. 2008). The virus-infected poison hemlock plants were stunted and showed chlorotic line pattern symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ApVY, refer to Apium graveolens.

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634

Conium maculatum (Hemlock or poison hemlock)

Carrot thin leaf virus Taxonomic position Genus: Potyvirus

(CTLV)

Family: Potyviridae

CTLV was detected in plants of Conium maculatum from Washington, USA (Howell and Mink 1981). The virus-infected poison hemlock plants exhibit symptomless infections. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CTLV, refer to Daucus carota.

Celery mosaic virus

(CeMV)

Synonyms Western celery mosaic virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

CeMV infection in plants of Conium maculatum was reported from California (Sutabutra and Campbell 1971; Howell and Mink 1981). The virus-infected poison hemlock plants exhibit systemic mottle and malformation symptoms or may be symptomless. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CeMV, refer to Apium graveolens.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Conium maculatum was reported from field surveys in New Zealand (Fletcher 2001). The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Conium maculatum was reported from Canada (Parrella et al. 2003). The virus-infected poison hemlock plants were symptomless. The virus is transmitted by a thrips vector, Frankliniella occidentalis, in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Convolvulus arvensis (Field bindweed)

635

References Eastwell KC, Glass JR, Seymour LM, Druffel KJ (2008) First report of infection of poison hemlock and celery by Apium virus Y in Washington State. Plant Dis 92:1710 Fletcher JD (2001) New hosts of Alfalfa mosaic virus, Cucumber mosaic virus, Potato virus Y, Soybean dwarf virus, and Tomato spotted wilt virus in New Zealand. N Z J Crop Hortic Sci 29:213–217 Howell WE, Mink GI (1981) Viruses isolated from wild carrot and poison hemlock. Plant Dis 65:277–279 Moran J, van Rijswijk B, Traicevski V, Kitajima EW, Mackenzie AM, Gibbs AJ (2002) Potyviruses, novel and known, in cultivated and wild species of the family Apiaceae in Australia. Arch Virol 147:1855–1967 Parrella G, Gognalons P, Gebre-Selassi K, Vovlas C, Marchoux G (2003) An update of the host range of Tomato spotted wilt virus. J Plant Pathol 85(4, special issue):227–264 Sutabutra T, Campbell RN (1971) Strains of Celery mosaic virus from parsley and poison hemlock in California. Plant Dis Reptr 55:328–332

Convolvulus arvensis (Field bindweed) Family: Convolvulaceae

Weed host

Alfalfa mosaic virus

(AMV)

Taxonomic position Genus: Alfamovirus

Family: Bromoviridae

AMV infection in plants of Convolvulus arvensis was reported from Saudi Arabia (Al-Shahwan et al. 2017). The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Beet curly top virus Taxonomic position Genus: Curtovirus

(BCTV)

Family: Geminiviridae

BCTV infection in plants of Convolvulus arvensis was reported from California, USA (Creamer et al. 1996). The virus is transmitted by a leafhopper vector, Circulifer tenellus, in a persistent (circulative, non-propagative) manner. The virus is not transmitted by mechanical sap-inoculation. For more details of BCTV, refer to Beta vulgaris.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Convolvulus arvensis was reported from Bulgaria and Southern New Mexico, USA (Dikova 1989; Rodriguez-Alvarado et al. 2002). The virus-infected field bindweed

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636

Convolvulus arvensis (Field bindweed)

plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Cucumber vein yellowing virus Taxonomic position Genus: Ipomovirus

(CVYV)

Family: Potyviridae

CVYV infection in plants of Convolvulus arvensis was reported from Spain (Janssen et al. 2002). The virus-infected field bindweed plants exhibit vein-clearing and leaf chlorosis symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CVYV, refer to Cucumis sativus.

Pepino mosaic virus Taxonomic position Genus: Potexvirus

(PepMV)

Family: Alphaflexiviridae

PepMV infection in plants of Convolvulus arvensis was reported from Spain and Cyprus (Cordoba et al. 2004; Papayiannis et al. 2012). The virus is mechanically sap-transmissible. For more details of PepMV, refer to Solanum muricatum.

Pepper mottle virus Taxonomic position Genus: Potyvirus

(PepMoV)

Family: Potyviridae

PepMoV infection in plants of Convolvulus arvensis was reported from Southern New Mexico, USA (Rodriguez-Alvarado et al. 2002). The virus-infected field bindweed plants exhibit symptoms of mosaic and distortion of foliage. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants. For more details of PepMoV, refer to Capsicum annuum.

Plum pox virus Taxonomic position Genus: Potyvirus

(PPV)

Family: Potyviridae

PPV infection in plants of Convolvulus arvensis was reported from Slovenia and Bulgaria (Virscek Marn et al. 2004; Milusheva and Rankova 2006). The virus-infected field bindweed plants exhibit pale or yellow-green ringspots or mottling on leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PPV, refer to Prunus domestica.

Convolvulus arvensis (Field bindweed)

637

Prunus necrotic ringspot virus Taxonomic position Genus: Ilarvirus

(PNRSV)

Family: Bromoviridae

PNRSV-infected plants of Convolvulus arvensis were reported from Iran (Sabaghian et al. 2013). The virusinfected field bindweed plants exhibit line patterns, mottle, and marginal necrosis symptoms. The virus is transmitted by the thrips vectors, and the virus present in/on pollen enters into the host through injuries caused by thrips while feeding. The virus is also transmissible by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants. For more details of PNRSV, refer to Prunus persica.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in plants of Convolvulus arvensis was reported from Bulgaria (Dikova 2006). The virusinfected field bindweed plants exhibit symptoms of yellow-orange-colored stems and fruits. The virus is transmitted by nematode vectors and also by mechanical sap-inoculation to a large number of host plants. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Convolvulus arvensis was reported from Spain and Cyprus (Jorda et al. 2001; Papayiannis et al. 2011). The virus-infected field bindweed plants exhibit symptomless infections. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not transmissible by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

References Al-Shahwan IM, Abdalla OA, Al-Saleh MA, Amer MA (2017) Detection of new viruses in alfalfa, weeds and cultivated plants growing adjacent to alfalfa fields in Saudi Arabia. Saudi J Biol Sci 24(6):1336–1343 Cordoba MC, Martinez-Priego LI, Jorda C (2004) New natural hosts of Pepino mosaic virus in Spain. Plant Dis 88:906 Creamer R, Lugue-Williams M, Howo M (1996) Epidemiology and incidence of Beet curly top geminivirus in naturally infected weed hosts. Plant Dis 80:533–535 Dikova B (1989) Wild-growing hosts of the Cucumber mosaic virus (Abstract). Rasteniev’dni Nauki 26(7):57–64 Dikova B (2006) Establishment of Tobacco rattle virus (TRV) in weeds and Cuscuta. Biotechnol Biotechnol Equip 20:42–48 Janssen D, Ruiz L, Velasco L, Segundo E, Cuadrado IM (2002) Non-cucurbitaceous weed species shown to be natural hosts of Cucumber vein yellowing virus in south eastern Spain. Plant Pathol 51:797 Jorda C, Font I, Martinez P, Juarez M, Ortega A, Lacasa A (2001) Current status and new natural hosts of Tomato yellow leaf curl virus (TYLCV) in Spain. Plant Dis 85:445

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Conyza spp.

Milusheva S, Rankova Z (2006) Serological identification of Plum pox virus in some economic important weeds. Selskostopanska Nauka (Agric Sci) 39:38–41 Papayiannis L, Katis NI, Idris AM, Brown JK (2011) Identification of weed hosts of Tomato yellow leaf curl virus in Cyprus. Plant Dis 95:120–125 Papayiannis LC, Kokkinos CD, Alfafo-Fernandez A (2012) Detection, characterization and host range studies of Pepino mosaic virus in Cyprus. Eur J Plant Pathol 132:1–7 Rodriguez-Alvarado G, Fernandez-Pavia S, Creamer R, Liddell C (2002) Pepper mottle virus causing disease in Chile peppers in southern New Mexico. Plant Dis 86:603–605 Sabaghian S, Rakhshandehroo F, Rezaee S (2013) First report of Prunus necrotic ringspot virus infecting bindweed in Iran. J Plant Pathol 95:667 Virscek Marn M, Mavric I, Urbancic-Zemljic M, Skerlavaj V (2004) Detection of Plum pox virus Potyvirus in weeds. Acta Hortic 657:251–254

Conyza spp. Synonyms Erigeron spp. Family: Asteraceae

Weed host

Lettuce mosaic virus Taxonomic position Genus: Potyvirus

(LMV)

Family: Potyviridae

LMV infection in plants of Erigeron bonariensis was reported from Brazil (Chaves et al. 2003). The virusinfected erigeron plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a nonpersistent manner, and also by mechanical sap-inoculation. For more details of LMV, refer to Lactuca sativa.

Pepino mosaic virus Taxonomic position Genus: Potexvirus

(PepMV)

Family: Alphaflexiviridae

PepMV infection in plants of Conyza albida was reported from Spain (Cordoba et al. 2004). The virus is mechanically sap-transmissible. For more details of PepMV, refer to Solanum muricatum.

Potato spindle tuber viroid Taxonomic position Genus: Pospiviroid

(PSTVd)

Family: Pospiviroidae

PSTVd infection in plants of Conyza bonariensis was reported from Australia (Mackie et al. 2016). The viroid-infected conyza plants exhibit symptoms of stunting, leaf chlorosis, and/or reduced leaf size. The viroid is mechanically sap-transmissible. For more details of PSTVd, refer to Solanum tuberosum.

Conyza spp.

639

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Conyza bonariensis was reported from Australia (Sharman et al. 2015). The virusinfected conyza plants exhibit leaf narrowing, deformation, and stunting symptoms. The virus is transmitted by the thrips vectors, and the virus present in/on the pollen enters into the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible. The virus is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato bushy stunt virus Taxonomic position Genus: Tombusvirus

(TBSV)

Family: Tombusviridae

TBSV infection in plants of Erigeron Canadensis was reported from Yugoslavia (Grbelja et al. 1988). The virus-infected erigeron plants exhibit mosaic symptoms. The virus is transmissible by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants. For more details of TBSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Erigeron canadensis was reported from Italy (Grieco et al. 2000). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sapinoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Conyza sumatrensis was reported from Spain and Cyprus (Jorda et al. 2001; Papayiannis et al. 2011). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not transmissible by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

Youcai mosaic virus Taxonomic position Genus: Tobamovirus

(YoMV)

Family: Virgaviridae

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Coprosma repens (Mirror bush)

YoMV infection in plants of Erigeron annuus was reported from South Korea (Park et al. 2016). The virus-infected erigeron plants exhibit mild mosaic and leaf malformation symptoms. There is no known vector for this virus. The virus is mechanically sap-transmissible and also transmissible through contact between plants. For more details of YoMV, refer to Brassica napus.

References Chaves ALR, Braun MR, Eiras M, Colariccio A, Galleti SR (2003) Erigeron bonariensis: an alternative host of Lettuce mosaic virus in Brazil. Fitopatol Bras 28:307–311 Cordoba MC, Martinez-Priego LI, Jorda C (2004) New natural hosts of Pepino mosaic virus in Spain. Plant Dis 88:906 Grbelja J, Eric Ž, Jeknik Z (1988) Erigeron canadensis L. – a potential source of infection of cultivated plants by Tomato bushy stunt virus. Fragm Herbologica Jugoslavia 17:95–99 Grieco PD, Conte D, Munno I, Nuzzaci M, de Stradis A (2000) Tomato spotted wilt virus (TSWV) on weeds and wild plants in Metapontino, Basilicata. Inf Fitopatol 50:43–46 Jorda C, Font I, Martinez P, Juarez M, Ortega A, Lacasa A (2001) Current status and new natural hosts of Tomato yellow leaf curl virus (TYLCV) in Spain. Plant Dis 85:445 Mackie AE, Rodoni BC, Barbetti MJ, McKirdy SJ, Jones RAC (2016) Potato spindle tuber viroid: alternative host reservoirs and strain found in a remote subtropical irrigation area. Eur J Plant Pathol 145:433–446 Papayiannis L, Katis NI, Idris AM, Brown JK (2011) Identification of weed hosts of Tomato yellow leaf curl virus in Cyprus. Plant Dis 95:120–125 Park CY, Lee M-A, Lee S-H, Kim J-S, Kim H-G (2016) First report of Youcai mosaic virus in daisy fleabane (Erigeron annuus). Plant Dis 100:1250 Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct Tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207

Coprosma repens (Mirror bush) Family: Rubiaceae

Ornamental

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Coprosma repens was reported from Italy (Polizzi and Bellardi 2007). The virus-infected mirror bush plants exhibit symptoms of irregular or semicircular necrotic spots, sometimes in concentric rings on leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Polizzi G, Bellardi MG (2007) First report of Tomato spotted wilt virus on Coprosma repens (mirror bush) in Italy. Plant Dis 91:1362

Corchorus spp. (Corchorus Capsularis, C. hirtus, C. olitorius, C. trilocularis) (Jute)

641

Corchoropsis timentosa Family: Malvaceae

Weed host

Papaya leaf curl China virus Taxonomic position Genus: Begomovirus

(PaLCuCNV)

Family: Geminiviridae

PaLCuCNV infection in plants of Corchoropsis timentosa was reported from China (Huang and Zhou 2006). The virus-infected Corchoropsis plants exhibit yellow vein symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of PaLCuCNV, refer to Carica papaya.

References Huang JF, Zhou XP (2006) First report of Papaya leaf curl China virus infecting Corchoropsis timentosa in China. Plant Pathol 55:291

Corchorus spp. (Corchorus Capsularis, C. hirtus, C. olitorius, C. trilocularis) (Jute) Family: Malvaceae

Fiber crop

Corchorus golden mosaic virus Taxonomic position Genus: Begomovirus

(CoGMV)

Family: Geminiviridae

Geographical distribution CoGMV infection in plants of Corchorus capsularis was reported from Vietnam, India, and Bangladesh (Ahmed 1978; Ha et al. 2008; Ghosh et al. 2008, 2011, 2012; Hasan et al. 2015). Symptoms and host(s) The virus-infected jute plants exhibit symptoms such as small yellow flecks on the lamina during the initial infection stage which gradually increase in size to form green and chlorotic intermingled patches producing a yellow mosaic appearance. Transmission The transmission of CoGMV has not been investigated. It is likely that, in common with other begomoviruses, CoGMV is transmitted by the whitefly vector Bemisia tabaci in a circulative, nonpropagative manner (Ahmed 1978).

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Corchorus spp. (Corchorus Capsularis, C. hirtus, C. olitorius, C. trilocularis) (Jute)

Virion properties and genome The structure of the virions of CoGMV has not been investigated. In common with all geminiviruses, the virions of CoGMV are likely to be geminated (twinned quasi-icosahedra). Although native to the Old World, CoGMV and Corchorus yellow vein virus are examples of begomoviruses having genomic features typical of begomoviruses native to the New World (Ha et al. 2008). The genome of CoGMV contained two circular, single-stranded DNA components. DNA-A contains 2677 nt (DQ641688 = NC_009644; FJ463902) and DNA-B of 2649 nt (DQ641689 = NC_009646; FJ463901) (Briddon 2001; Ha et al. 2008; Hasan and Sano 2014; Brown et al. 2015; Hasan et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the New World, the DNA A component of CoGMV encodes five genes, one in the virion sense and four in the complementary sense. The AV2 gene common to the Old World begomoviruses is absent in CoGMV. The DNA B encodes one gene in each orientation. The expression and function of these genes have not been investigated for CoGMV.

Corchorus yellow spot virus Taxonomic position Genus: Begomovirus

(CoYSV)

Family: Geminiviridae

Geographical distribution CoYSV infection in plants of Corchorus siliquosus was reported from Mexico (Hernandez-Zepeda et al. 2007). The virus was previously known as Corchorus yellow vein Yucatan virus. Symptoms and host(s) The virus-infected jute plants exhibit yellow mosaic and yellow vein symptoms. Transmission The transmission of the CoYSV has not been investigated. It is likely that, in common with other begomoviruses, CoYSV is transmitted by the whitefly vector Bemisia tabaci in a circulative, nonpropagative manner. Virion properties and genome The structure of the virions of CoYSV has not been investigated. In common with all geminiviruses, the virions of CoYSV are likely geminate (twinned quasi-icosahedra). CoYSV is typical of the majority of begomoviruses native to the New World with a bipartite genome consisting of two circular, single-stranded DNA components. DNA-A contains 2606 nt (DQ875868 = NC_008492) and DNA-B of 2589 nt (DQ875869 = NC_008493) (Briddon 2001; Hernandez-Zepeda et al. 2007; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the New World, the DNA-A component of CoYSV encodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes has not been investigated for CoYSV.

Corchorus yellow vein mosaic virus Taxonomic position Genus: Begomovirus

(CoYVMV)

Family: Geminiviridae

Corchorus spp. (Corchorus Capsularis, C. hirtus, C. olitorius, C. trilocularis) (Jute)

643

Geographical distribution CoYVMV infection in plants of Corchorus olitorius was reported from India (Archana and Malathi unpublished - KC223600). Symptoms and host(s) The virus-infected jute plants exhibit yellow vein mosaic symptoms. Transmission The transmission of CoYVMV has not been investigated. It is likely that, in common with other begomoviruses, CoYVMV is transmitted by the whitefly vector Bemisia tabaci in a circulative, nonpropagative manner. Virion properties and genome The structure of the virions of CoYVMV has not been investigated. In common with all geminiviruses, the virions of CoYVMV are likely geminate (twinned quasi-icosahedra). With only two isolates of CoYVMV having been characterized, the precise nature of the genome (whether monopartite, monopartite betasatellite-associated, or bipartite) is unclear. The characterized genome component of CoYVMV contained a single stranded circular DNA of ~2743 nt (KC223600 = NC_020473; KC196077) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). The expression and function of the genes have not been investigated.

Corchorus yellow vein virus

(CoYVV)

Synonyms Corchorus yellow vein Vietnam virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution CoYVV infection in plants of Corchorus capsularis was reported from Vietnam and China (Ha et al. 2006; Lin et al. 2016). Symptoms and host(s) The virus-infected jute plants exhibit vein yellowing symptoms. Transmission Insect transmission of CoYVV has not been demonstrated. However, in common with all other begomoviruses, CoYVV is likely transmitted by the whitefly vector of begomoviruses, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of CoYVV has not been investigated. In common with all geminiviruses, the virions of CoYVV are likely geminate (twinned quasi-icosahedra). Although native to the Old World, the CoYVV genome has features typical of begomoviruses native to the New World (Ha et al. 2006). The genome contains two circular, single-stranded DNA components. DNA-A consists of 2724 nt (AY727903 = NC_006358) and DNA-B is 2691 nt

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Corchorus spp. (Corchorus Capsularis, C. hirtus, C. olitorius, C. trilocularis) (Jute)

(AY727904 = NC_006359) (Briddon 2001; Ha et al. 2006; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the New World, the DNA-A component of CoYVV encodes five genes (lacking AV2 gene), one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The function of these genes has not been investigated for CoYVV.

Groundnut bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(GBNV)

Family: Tospoviridae

GBNV-infected plants of Corchorus spp. were reported from India (Sivaprasad et al. 2011). The virusinfected jute plants showed mosaic, chlorotic, and necrotic lesions, which occurred on young leaves and stems, and ultimately resulted in plant death (Sivaprasad et al. 2011). The virus is transmitted by a thrips vector, Thrips palmi, in a persistent-propagative manner, and also by mechanical sap-inoculation to a wide range of host plants. For more details of GBNV, refer to Arachis hypogaea.

Okra mosaic virus Taxonomic position Genus: Tymovirus

(OkMV)

Family: Tymoviridae

OkMV infection in plants of Corchorus olitorius was reported from the Ivory Coast (Givord 1978). The virus-infected jute plants exhibit vein chlorosis and vein-banding symptoms. The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sap-inoculation, and by grafting. For more details of OkMV, refer to Abelmoschus esculentus.

Potato leafroll virus Taxonomic position Genus: Polerovirus

(PLRV)

Family: Luteoviridae

PLRV infection in plants of Corchorus olitorius was reported from India (Biswas et al. 2014). The virus-infected jute plants exhibit symptoms of stunted growth and short height. Mainly the upper leaves are elongated with curling and coiling of the lamina. Puckering and shoestring effect were also noticed. Petioles and stipules of the affected leaves were exceptionally longer. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical sapinoculation. For more details of PLRV, refer to Solanum tuberosum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Corchorus trilocularis was reported from Australia (Sharman et al. 2015). The virus is transmitted by thrips vectors, and the virus present in/on the pollen entering into the host

Corchorus spp. (Corchorus Capsularis, C. hirtus, C. olitorius, C. trilocularis) (Jute)

645

through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Corchorus capsularis and C. olitorius was reported from Saudi Arabia (Sohrab 2016). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative nonpropagative manner. The virus is transmissible by grafting but not transmissible by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

References Ahmed M (1978) A whitefly vectored yellow mosaic of jute. FAO Plant Protect Bull 26:169–171 Biswas C, Dey P, Mitra S, Bera A, Satpathy S, Karmakar PG (2014) First report of Potato leaf roll virus (PLRV) naturally occurring on Jute (Corchorus olitorius) in India. Plant Dis 98:1592 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Ghosh R, Paul S, Das S, Palit P, Acharyya S, Das A, Mir JI, Ghosk SK, Roy A (2008) Molecular evidence for existence of a New World Begomovirus associated with yellow mosaic disease of Corchorus capsularis in India. Aust Plant Dis Notes 3:59–62 Ghosh R, Palit P, Ghosh SK, Roy A (2011) A new world virus alters biochemical profiling of jute plants (Corchorus capsularis) upon infection. Int J Sci Nat 2(4):883–885 Ghosh R, Palit P, Paul S, Ghosh SK, Roy A (2012) Detection of Corchorus golden mosaic virus associated with yellow mosaic disease of jute (Corchorus capsularis). Indian J Virol 23:70–74 Givord L (1978) Alternative hosts of Okra mosaic virus near plantings of okra in Southern Ivory Coast. Plant Dis Reptr 62:412–416 Ha C, Coombs S, Revill P, Harding R, Vu M, Dale J (2006) Corchorus yellow vein virus, a New World geminivirus from the old world. J Gen Virol 87:997–1003 Ha C, Coombs S, Revill P, Harding R, Vu M, Dale J (2008) Molecular characterization of begomoviruses and DNA satellites from Vietnam: additional evidence that the New World geminiviruses were present in the Old World prior to continental separation. J Gen Virol 89:312–326 Hasan MM, Sano Y (2014) Genomic variability of Corchorus golden mosaic virus originating from Bangladesh. Int J Phytopathol 3:81–88 Hasan MM, Meah MB, Ali MA, Okazaki K, Sano Y (2015) Characterization and confirmation of Corchorus golden mosaic virus associated with Jute in Bangladesh. J Plant Pathol Microb 6:256 Hernandez-Zepeda C, Idris AM, Carnevali G, Brown JK, Moreno-Valenzuela OA (2007) Molecular characterization and phylogenetic relationships of two new bipartite Begomovirus infecting malvaceous plants in Yucatan, Mexico. Virus Genes 35:369–377 Lin WZ, Xiong GH, Qiu P, Wu KC, Du ZG, Zhang J, Wu ZJ (2016) First report of the occurrence of Corchorus yellow vein Vietnam virus on Jute in China. Plant Dis 100:2176 Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct Tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207 Sivaprasad Y, Bhaskara Reddy BV, Naresh Kumar CVM, Raja Reddy K, Sai Gopal DVR (2011) Jute (Corchorus capsularis): a new host of Peanut bud necrosis virus. New Dis Rep 23:33 Sohrab SS (2016) The role of corchorus in spreading of Tomato yellow leaf curl virus on tomato in Jeddah, Saudi Arabia. Virus Dis 27:19–26 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

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646

Cordia crenata

Cordia crenata Family: Boraginaceae

Ornamental

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV was reported to infect Cordia crenata in Iran (Parrella et al. 2003; Ghotbi et al. 2005). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Ghotbi T, Sharaeen N, Winter S (2005) Occurrence of tospoviruses in ornamental and weed species in Markazi and Tehran provinces in Iran. Plant Dis 89:425–429 Parrella G, Gognalons P, Gebre-Selassi K, Vovlas C, Marchoux G (2003) An update of the host range of Tomato spotted wilt virus. J Plant Pathol 85(4, special issue):227–264

Cordyline fruticosa (Ti plant, Good luck plant) Synonyms Cordyline terminalis Family: Asparagaceae

Cordyline virus 1 Taxonomic position Genus: Velarivirus

Ornamental

(CoV-1)

Family: Closteroviridae

Geographical distribution CoV-1 infection in plants of Cordyline fruticosa was reported from Hawaii (Melzer et al. 2011, 2013a, b). Symptoms and host(s) The virus-infected good luck plants exhibit ringspot symptoms.

Cordyline fruticosa (Ti plant, Good luck plant)

647

Transmission The virus is not mechanically transmissible and has no known vector; an insect vector is suspected. Virion properties and genome The virions are non-enveloped, filamentous particles of about 1500–1700 nm in length. The genome is a linear positive-sense single-stranded RNA of 16,833 nt (HM588723) which encodes 9 ORFs (Martelli et al. 2012).

Cordyline virus 2 Taxonomic position Genus: Velarivirus

(CoV-2)

Family: Closteroviridae

Geographical distribution CoV-2 infection in plants of Cordyline fruticosa was reported from Hawaii (Melzer et al. 2013a, b). Symptoms and host(s) The virus-infected good luck plants exhibit ringspot symptoms. Transmission No vector identified but likely to be insect-transmitted.

Virion properties and genome The virions are non-enveloped, filaments particles of about 1500–1700 nm in length. The genome is a linear positive-sense single-stranded RNA of >15 kb; a close to full sequence of c.15.1 kb is available (JQ599282) (Melzer et al. 2013b).

Cordyline virus 3 Taxonomic position Genus: Velarivirus

(CoV-3)

Family: Closteroviridae

Geographical distribution CoV-3 infection in plants of Cordyline fruticosa was reported from Hawaii (Melzer et al. 2013a, b).

Symptoms and host(s) The virus-infected good luck plants exhibit ringspot symptoms.

Transmission No vector identified but likely to be insect-transmitted.

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648

Cordyline fruticosa (Ti plant, Good luck plant)

Virion properties and genome The virions are non-enveloped, filaments particles of about 1500–1700 nm in length. The genome is a linear positive-sense single-stranded RNA of >16 kb; a close to full sequence of c.16.3 kb is available (JQ599283) (Melzer et al. 2013b).

Cordyline virus 4 Taxonomic position Genus: Velarivirus

(CoV-4)

Family: Closteroviridae

Geographical distribution CoV-4 infection in plants of Cordyline fruticosa was reported from Hawaii (Melzer et al. 2013a, b). Symptoms and host(s) The virus-infected good luck plants exhibit ringspot symptoms. Transmission No vector identified but likely to be insect-transmitted. Virion properties and genome The virions are non-enveloped, filaments particles of about 1500–1700 nm in length. The genome is a linear positive-sense single-stranded RNA of >14 kb; a close to full sequence of c.14.6 kb is available (JQ599284) (Melzer et al. 2013b).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Cordyline terminalis was reported from Italy (Roggero et al. 1999). The virus-infected good luck plants show chlorotic-necrotic ringspots and leaf malformation symptoms. The virus is transmitted by a thrips vector, Frankliniella occidentalis, in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Cordyline terminalis was reported from Hawaii (Cho et al. 1987). The virus-infected good luck plants exhibit chlorotic or necrotic spots on the leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Coriandrum sativum (Coriander or Cilantro)

649

References Cho JJ, Mau RFL, Mitchell WC, Gonsalves D, Yudin LS (1987) Host list of plants susceptible to Tomato spotted wilt virus (TSWV), Research Extension Series; RES-078. University of Hawaii. 10 p, Honolulu Martelli GP, Abou Ghanem-Sabanadzovic N, Agranovsky AA, Al Rwahnih M, Dolja VV, Dovas CI, Fuchs M, Gugerli P, Hu JS, Jelkmann W, Katis NI, Maliogka VI, Melzer MJ, Menzel W, Minafra A, Rott ME, Rowhani A, Sabanadzovic S, Saldarelli P (2012) Taxonomic revision of the family Closteroviridae with special reference to the grapevine leafroll – associated members of the genus Ampelovirus and the putative species unassigned to the family. J Plant Pathol 94:7–19 Melzer MJ, Sehter DM, Borth WB, Mersino EF, Hu JS (2011) An assemblage of Closteroviruses infects Hawaiian ti (Cordyline fruticosa L.). Virus Genes 42:254–260 Melzer M, Ayin C, Sugano J, Uchida J, Kawate M, Borth W, Hu J (2013a) Differentiation and distribution of cordyline viruses 1–4 in Hawaiian ti plants (Cordyline fruticosa L.). Viruses 5:1655–1663 Melzer MJ, Sugano JS, Uchida JY, Kawate MK, Borth WB, Sether DM, Hu JS (2013b) Molecular characterization of closteroviruses infecting Cordyline fruticosa (L.) in Hawaii. Front Microb 4:39 Roggero P, Ciuffo M, Dellavalle G, Gotta P, Gallo S, Peters D (1999) Additional ornamental species as hosts of Impatiens necrotic spot tospovirus in Italy. Plant Dis 83:967

Coreopsis spp. (C. auriculata, C. lanceolata) (Tickseed) Family: Asteraceae

Weed host

Lettuce mosaic virus Taxonomic position Genus: Potyvirus

(LMV)

Family: Potyviridae

LMV infection in plants of Coreopsis auriculata was reported from the USA (Naidu et al. 2008). The virus-infected tickseed plants exhibit symptoms of chlorotic spots and rings. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of LMV, refer to Lactuca sativa.

References Naidu RA, Karthikeyan G, Jarugula S, Nelson M, Morrell A (2008) First report of the natural infection of Coreopsis auriculata ‘Nana’ with Lettuce mosaic virus in the United States of America. Plant Dis 92:486

Coriandrum sativum (Coriander or Cilantro) Family: Apiaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Spice

(AMV)

Family: Bromoviridae

C

650

Coriandrum sativum (Coriander or Cilantro)

AMV infection in plants of Coriandrum sativum was reported from New Zealand and Yemen (Alhubaishi et al. 1987; Fletcher 1989). The virus-infected coriander plants exhibit symptoms of interveinal mottling, chlorosis of young leaves, bunching, and distortion of growing points. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Apium virus Y

(ApVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

ApVY infection in plants of Coriandrum sativum was reported from California, USA (Tian et al. 2008). The virus-infected coriander plants exhibit symptoms of mosaic, vein-clearing, and stunting. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ApVY, refer to Apium graveolens.

Celery yellow mosaic virus

(CeYMV)

Taxonomic position CeYMV is a tentative member of the genus Potyvirus and family Potyviridae. CeYMV infection in plants of Coriandrum sativum was reported from Egypt (Abo El-Ela et al. 2005). The virus-infected coriander plants exhibit yellow and mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CeYMV, refer to Apium graveolens.

Clover yellow vein virus Taxonomic position Genus: Potyvirus

(ClYVV)

Family: Potyviridae

ClYVV infection in plants of Coriandrum sativum was reported from Canada (Singh and Lopez-Abella 1971). The virus-infected coriander plants exhibit symptoms of vein-clearing which resulted in yellownet appearance. In addition, leaves were severely malformed with cupping and plants were severely stunted. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ClYVV, refer to Trifolium spp.

Coriander feathery red vein virus

(CFRVV)

Taxonomic position CFRVV is a tentative member of the family Rhabdoviridae. Geographical distribution CFRVV was first reported in Coriandrum sativum plants from the USA by Misari and Sylvester (1983). The virus spreads in the USA (California).

Coriandrum sativum (Coriander or Cilantro)

651

Symptoms and host(s) The virus-infected coriander plants exhibit red vein-banding symptoms. Transmission The virus is transmitted by aphid vectors, Hyadaphis foeniculi and Myzus persicae, in a persistent manner. The virus is not transmitted by Aphis fabae, A. helianthi heraciella, Acyrthosiphon (Aulacorthum) solani, Cavariella aegopodii, Chaetosiphon thomasi jacobi, Dysaphis apiifolia, Hyadaphis erysimi, Hyperomyzus lactucae, Myzus ornatus, and Aulacorthum circumflexum. The virus multiplies in the vector and is transmitted congenitally to the progeny of the vector. The virus is transmissible by mechanical sap-inoculation, and not transmitted by seed. Virion properties and genome The virions are rhabdo- or bullet-shaped, enveloped with a clear modal length of 216 nm (in sections) and 75 nm wide (in sections).

Groundnut ringspot orthotospovirus Taxonomic position Genus: Orthotospovirus

(GRSV)

Family: Tospoviridae

GRSV infection in plants of Coriandrum sativum was reported from Brazil (Lima et al. 1999). The virus-infected coriander plants exhibit symptoms of stunting, chlorotic ringspotting, necrosis, and malformation of apical leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of GRSV, refer to Arachis hypogaea.

Vanilla distortion mosaic virus Taxonomic position Genus: Potyvirus

(VDMV)

Family: Potyviridae

VDMV infection in plants of Coriandrum sativum was reported from India (Adams et al. 2014). The virus-infected coriander plants exhibit symptoms of chlorosis, bronzing, and necrosis on petioles and stems. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of VDMV, refer to Vanilla spp.

References Abo El-Ela AA, Amer MA, Abo El-Abbas F (2005) Celery yellow mosaic Potyvirus affecting umberlliferae plants in Egypt. Egypt J Virol 2(1):269–282 Adams IP, Rai S, Deka M, Harju V, Hodges T, Hayward G, Skelton A, Fox A, Boonham N (2014) Genome sequence of Vanilla distortion mosaic virus infecting Coriandrum sativum. Arch Virol 159:3463–3465 Alhubaishi AA, Walkey DGA, Webb MJW, Bolland CJ, Cook AA (1987) A survey of horticultural plant virus diseases in the Yemen Arab Republic. FAO Plant Protect Bull 35:135–143 Fletcher JD (1989) Additional hosts of Alfalfa mosaic virus, Cucumber mosaic virus and Tobacco mosaic virus in New Zealand. N Z J Crop Hortic Sci 17:361–362 Lima MF, de Avila AC, da Wanderley LJG Jr, Nagata T, da Gama LJW (1999) Coriander: a new natural host of groundnut ringspot in Brazil. Plant Dis 83:878

C

652

Cornus florida (Flowering dogwood)

Misari SM, Sylvester ES (1983) Coriander feathery red-vein virus, a propagative plant Rhabdovirus, and its transmission by the aphid Hyadaphis foeniculi Passerini. Hilgardia 51:1–38 Singh RP, Lopez-Abella D (1971) Natural infection of coriander plants by a strain of Clover yellow vein virus. Phytopathology 61:333–334 Tian T, Liu H-Y, Koike ST (2008) First report of Apium virus Y on cilantro, celery, and parsley in California. Plant Dis 92:1254

Cornus florida (Flowering dogwood) Family: Cornaceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

Family: Secoviridae

ArMV infection in plants of Cornus florida was reported from South Carolina (Barnett and Baxter 1976; Barnett 1981). The virus-infected flowering dogwood plants exhibit white or chlorotic mosaic symptoms. The virus is transmitted by nematode vectors (Xiphinema spp.) in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Broad bean wilt virus Taxonomic position Genus: Fabavirus

(BBWV)

Family: Secoviridae

BBWV (but without specifying either of the currently recognized species BBWV-1 or BBWV-2) infection in plants of Cornus florida was reported from South Carolina (Scott and Barnett 1984). The virus-infected flowering dogwood plants exhibit faint yellow mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BBWV, refer to Vicia faba.

Cherry leaf roll virus Taxonomic position Genus: Nepovirus

(CLRV)

Family: Secoviridae

CLRV was detected from trees of Cornus florida in the USA including Florida (Reddick et al. 1979; von Bargen et al. 2009). The virus-infected flowering dogwood plants occasionally show ringspots but typically do not show apparent symptoms (Waterworth and Lawson 1973). The virus is transmitted by nematode vectors in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and also by grafting. For more details of CLRV, refer to Prunus avium.

Cornus florida (Flowering dogwood)

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

653

(TRSV)

Family: Secoviridae

TRSV was reported from Cornus florida in the USA (Waterworth and Povish 1972; Reddick et al. 1979). The virus-infected flowering dogwood plants do not exhibit any ringspot symptoms. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato bushy stunt virus Taxonomic position Genus: Tombusvirus

(TBSV)

Family: Tombusviridae

TBSV infection in plants of Cornus florida was reported from the UK (Brunt and Stace-Smith 1971). The virus-infected flowering dogwood plants exhibit yellow ring and line pattern symptoms. The virus is transmissible by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants. There is no known vector for this virus. For more details of TBSV, refer to Solanum lycopersicum.

Tomato mosaic virus Taxonomic position Genus: Tobamovirus

(ToMV)

Family: Virgaviridae

ToMV infection in plants of Cornus florida was reported from Tennessee (Reddick 1989). The virusinfected flowering dogwood plants exhibit systemic mosaic symptoms. No vector is involved in the spread of this virus. This virus is transmissible by mechanical sap-inoculation, transmissible by grafting, and also transmissible by contact between plants. For more details of ToMV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Cornus florida was reported from South Carolina (Reddick et al. 1979). The virus-infected flowering dogwood plants are typically symptomless. The virus is transmitted by nematode vectors in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and also by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

C

654

Corylus spp. (Hazelnut)

References Barnett OW (1981) Arabis mosaic virus from Cornus florida in South Carolina. In: Fifth international congress of virology, abstracts, Strasbourg, France. p 237 Barnett OW, Baxter LW (1976) Arabis mosaic virus from a wild dogwood in South Carolina. Proc Am Phytopathol Soc 3:249 Brunt AA, Stace-Smith R (1971) Tomato bushy stunt virus on dogwood. Annual Rept Glasshouse Crops Res Inst, Littlehampton, Sussex. pp 118–122 Reddick BB (1989) Isolation and partial characterization of a Tobamovirus from flowering dogwood in Tennessee. Plant Dis 73:174–176 Reddick BB, Barnett OW, Baxter LW (1979) Isolation of cherry leaf roll, tobacco ringspot and tomato ringspot viruses from dogwood trees in South Carolina. Plant Dis Reptr 63:529–532 Scott SW, Barnett OW (1984) Some properties of an isolate of Broad bean wilt virus from dogwood (Cornus florida). Plant Dis 68:983–985 von Bargen S, Grubits E, Jalkanen R, Büttner C (2009) Cherry leaf roll virus – an emerging virus in Finland? Silva Fenn 43:727–738 Waterworth HE, Lawson RH (1973) Purification, electron microscopy and serology of the dogwood ringspot strain of Cherry leaf roll virus. Phytopathology 63:141–146 Waterworth HE, Povish WR (1972) Tobacco ringspot virus from naturally infected dogwood, autumn crocus, and Forsythia. Plant Dis Reptr 56:336–337

Corylus spp. (Hazelnut) Family: Betulaceae

Trees/Shrubs

Apple mosaic virus

(ApMV)

Taxonomic position Genus: Ilarvirus

Family: Bromoviridae

ApMV infection in plants of Corylus spp. plants was reported from Turkey, Italy, France, Spain, the UK, Poland, and the USA (Postman and Mahlenbacher 1994; Aramburu and Rovira 1995, 1998, 2000; Piskornik et al. 2002; Arli Sokmen et al. 2004, 2005; Akbas et al. 2005; Kobylko et al. 2005; Akbas and Degirmenci 2009, 2010; Ertunc et al. 2014; Cieslinska and Valasevich 2016). The virus-infected hazelnut plants exhibit symptoms of yellow bands, spots, and oak leaf pattern on leaves. The upper leaves show systemic chlorosis and necrotic lines. No insect vector is known for this virus. The virus is transmissible by mechanical sap-inoculation. The virus is seed-transmitted through the seed of Corylus avellana up to 2% (Cameron and Thompson 1986). For more details of ApMV, refer to Malus domestica.

References Akbas B, Degirmenci K (2009) Incidence and natural spread of Apple mosaic virus on hazelnut in the west black sea coast of Turkey and its effect on yield. J Plant Pathol 91:767–771 Akbas B, Degirmenci K (2010) Simultaneous detection of Apple mosaic virus in cultivated hazelnuts by one-tube RT-PCR. Afr J Biotechnol 9:1753–1757 Akbas B, Ilhan D, Atlamaz A (2005) A preliminary survey of hazelnut (Corylus avellana L.) viruses in Turkey. Acta Hortic 686:359–466 Aramburu JM, Rovira M (1995) Effect of Apple mosaic virus on the growth and yield of “Negret” hazelnut. Acta Hortic 386:565–568 Aramburu J, Rovira M (1998) The effects of Apple mosaic ilarvirus (ApMV) on hazelnut (Corylus avellana L.). J Hortic Sci Biotechnol 73:97–101

Corypha utan (Buri palm)

655

Aramburu J, Rovira M (2000) Incidence and natural spread of Apple mosaic ilarvirus in hazel in north-east Spain. Plant Pathol 49:423–427 Arli Sokmen M, Sevik MA, Yilmaz MA (2004) Incidence of Apple mosaic virus infection in hazelnut (Corylus avellana L.) orchards of Samsun province. In: Proceedings of the first plant protection congress of Turkey, Samsun 2004. p 173 Arli Sokmen M, Kutluk Yilmaz ND, Mennan H, Sevik MA (2005) Natural weed hosts of Apple mosaic virus in hazelnut orchards in Turkey. J Plant Pathol 87:239–242 Cameron HR, Thompson M (1986) Seed transmission of Apple mosaic virus in hazelnut. Acta Hortic 193:131–132 Cieslinska M, Valasevich N (2016) Characterization of Apple mosaic virus isolates detected in hazelnut in Poland. J Plant Dis Prot 123:187–192 Ertunc F, Topkaya S, Sezer A (2014) Distribution and molecular detection of Apple mosaic virus in apple and hazelnut in Turkey. Afr J Biotechnol 13:3144–3149 Kobylko T, Nowak B, Urban A (2005) Incidence of Apple mosaic virus (ApMV) on hazelnut in south-east Poland. Folia Hortic 17:153–161 Piskornik Z, Kobyłko T, Nowak B (2002) Detection of Apple mosaic virus (ApMV) on hazelnut (Corylus sp.) in Poland. Phytopathol Pol 23:31–37 Postman JD, Mahlenbacher SA (1994) Apple mosaic virus in hazelnut germplasm. Acta Hortic 351:601–605

Corynocarpus laevigatus (Karaka or New Zealand laurel) Family: Corynocarpaceae

Trees/Shrubs

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Corynocarpus laevigatus was reported from New Zealand (Ashby 1977). The virus-infected karaka plants exhibit line-pattern and mottle symptoms on leaves. Symptoms tended to fade and disappear during summer. The virus is transmitted by the aphid vector, Myzus persicae, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

References Ashby JW (1977) Infection of karaka (Corynocarpus laevigatus JR and G Forst) by Cucumber mosaic virus. N Z J Agric Res 20:533–534

Corypha utan (Buri palm) Synonyms Corypha elata Family: Arecaceae

Palm tree, Commercial crop

C

656

Cosmos spp.

Coconut cadang-cadang viroid Taxonomic position Genus: Cocadviroid

(CCCVd)

Family: Pospiviroidae

CCCVd was reported from buri palm (Corypha elata) from in the Philippines (Randles et al. 1980). The viroid-infected buri palm plants show reduced frond size and produce large pale green to yellow spots on the third to fourth youngest opened frond. Four to five years after inoculation, the infected palms were usually stunted compared with uninoculated palms. Yellowing of leaflets was observed with defined spots or mottling of the older fronds. All infected palms showed mild or severe yellow-leaf spotting (Imperial et al. 1985). The viroid was successfully transmitted to seedlings of Areca catechu (betel nut palm), Corypha elata (buri palm), Adonidia merrillii (Manila palm), Elaeis guineensis (oil palm), Chrysalidocarpus lutescens (palmera), and Oreodoxa regia (royal palm). For more details of CCCVd, refer to Cocos nucifera.

References Imperial JS, Bautista RM, Randles JW (1985) Transmission of the Coconut cadang-cadang viroid to six species of palm by inoculation with nucleic acid extracts. Plant Pathol 34(3):391–401 Randles JW, Boccardo G, Imperial JS (1980) Detection of the cadang-cadang associated RNA in African oil palm and buri palm. Phytopathology 70(3):185–189

Cosmos spp. Family: Asteraceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

Family: Secoviridae

ArMV was reported affecting Cosmos spp. in the USA (Lawson and Hsu 2006). The virus is transmitted by nematode vectors (Xiphinema spp.) in a non-persistent manner, and also by mechanical sapinoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Beet curly top virus Taxonomic position Genus: Curtovirus

(BCTV)

Family: Geminiviridae

BCTV infection in plants of Cosmos bipinnatus was reported primarily from the Western USA (Smith 1972). The virus-infected cosmos plants exhibit shortened internodes toward the tips of the stems, with yellowing of secondary shoots and the apices of branches; leaves may be curled and twisted, with down-

Cotula australis (Annual buttonweed)

657

turned petioles. On secondary shoots, small chlorotic leaves may surround dwarfed flowers. The virus is transmitted by leafhopper vectors in a persistent (circulative, non-propagative) manner. The virus is not transmissible by mechanical sap-inoculation. For more details of BCTV, refer to Beta vulgaris.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

C Family: Bromoviridae

CMV infection in plants of Cosmos sulphureus was reported from Bangladesh (Parvin et al. 2007). The virus-infected cosmos plants show various combinations of mosaic, yellowing, shoe-stringing or curling of leaves, severe stunt, and leaf necrosis. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Cosmos bipinnatus and C. sulphureus has been reported by Parrella et al. (2003) and Cho et al. (1987). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Cho JJ, Mau RFL, Mitchell WC, Gonsalves D, Yudin LS (1987) Host list of plants susceptible to Tomato spotted wilt virus (TSWV). University of Hawaii, Honolulu 10 p. (Research Extension Series; RES-078) Lawson R, Hsu HT (2006) Quarantine viruses, viroids, and phytoplasmas that affect movement of ornamental plants. Acta Hortic 722:17–30 Parrella G, Gognalons P, Gebre-Selassi K, Vovlas C, Marchoux G (2003) An update of the host range of Tomato spotted wilt virus. J Plant Pathol 85(4):227–264 Parvin MS, Akanda AM, Rahman AHMA (2007) Summer cosmos – a host of Cucumber mosaic virus. J Agric Rural Dev 5:84–93 Smith KM (1972) A textbook of plant virus diseases, 3rd edn. Academic, New York

Cotula australis (Annual buttonweed) Family: Asteraceae

Weed host

Potato virus Y

(PVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

658

Cotyledon orbiculata (Pig’s ear)

PVY infection in plants of Cotula australis was reported from New Zealand (Fletcher 2001). The virusinfected plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts and by grafting. For more details of PVY, refer to Solanum tuberosum.

References Fletcher D (2001) New hosts of Alfalfa mosaic virus, Cucumber mosaic virus, Potato virus Y, Soybean dwarf virus, and Tomato spotted wilt virus in New Zealand. N Z J Crop Hortic Sci 29:213–217

Cotyledon orbiculata (Pig’s ear) Family: Crassulaceae

Medicinal

Cotyledon virus Y

(CotVY)

Taxonomic position CotVY is a tentative member of the genus Potyvirus and family Potyviridae Geographical distribution CotVY infection in plants of Cotyledon orbiculata was reported from Brazil (Duarte et al. 2014). Symptoms and host(s) The virus-infected pig’s ear plants exhibit foliar mosaic and distortion symptoms. Transmission The virus is transmitted by the aphid vector, Myzus persicae, and also by mechanical sap-inoculation. Virion properties and genome The virions are flexuous filaments. The genome is a single molecule of positive-sense single-stranded RNA presumed to be of 9.5–10 kb. A partial genome sequence of 1752 nt is available (JN572103) (Duarte et al. 2014; Wylie et al. 2017).

References Duarte LML, Alexandre MAV, Rivas EB, Galleti SR, Harakava R, Chaves ALR (2014) A new Potyvirus species in Cotyledon orbiculata in mixed infection with a Nucleorhabdovirus. J Plant Pathol 96:143–149 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Crambe spp. (C. abyssinica, C. glabrata, and C. hispanica)

659

Crambe spp. (C. abyssinica, C. glabrata, and C. hispanica) Family: Brassicaceae

Commercial crop

Beet western yellows virus Taxonomic Position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

BWYV infection in plants of Crambe abyssinica was reported from California, USA (Duffus 1975). The virus-infected plants exhibit stunting and interveinal reddening symptoms. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. For more details of BWYV, refer to Beta vulgaris.

Cauliflower mosaic virus Taxonomic Position Genus: Caulimovirus

(CaMV)

Family: Caulimoviridae

CaMV infection in plants of Crambe spp. was reported from Hungary (Horvath and Besada 1980). The virus is transmitted by aphid vectors in a semi-persistent manner, and also by mechanical sapinoculation. For more details of CaMV, refer to Brassica oleracea var. botrytis.

Croton yellow vein mosaic virus Taxonomic Position Genus: Begomovirus

(CroYVMV)

Family: Geminiviridae

CroYVMV infection in plants of Crambe spp. was reported from India (Kumar et al. 2017). The virusinfected plants exhibit typical leaf curling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of CroYVMV, refer to Codiaeum variegatum.

Radish mosaic virus Taxonomic Position Genus: Comovirus

(RaMV)

Family: Secoviridae

RaMV infection in plants of Crambe abyssinica was reported from Europe (Horvath et al. 1973). The virus is transmitted by beetle vectors and also by mechanical sap-inoculation, and by grafting. For more details of RaMV, refer to Raphanus sativus.

C

660

Crassocephalum crepidioides (Thickhead)

Tobacco mosaic virus

(TMV)

Taxonomic Position Genus: Tobamovirus

Family: Virgaviridae

TMV infection in plants of Crambe abyssinica was reported from the USA (Leppik 1973). There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Turnip yellow mosaic virus Taxonomic Position Genus: Tymovirus

(TYMV)

Family: Tymoviridae

TYMV infection in plants of Crambe abyssinica was reported from Europe (Horvath et al. 1973). The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sapinoculation, and by grafting. For more details of TYMV, refer to Brassica rapa.

References Duffus JE (1975) Effects of beet western yellows virus on crambe in the greenhouse. Plant Dis Rptr 59:886–888 Horvath J, Besada WH (1980) Herbaceous and Woody Crambe species as new artificial hosts of cauliflower mosaic virus. J Phytopathol 99:327 Horvath J, Juretić N, Miličič D (1973) Crambe abyssinica Hochst. Ex R. E. Frees as a new host plant for turnip yellow mosaic virus and radish mosaic virus. J Phytopathol 78:69–74 Kumar A, Bag MK, Singh R, Jailani AAK, Mandal B, Roy A (2017) Natural infection of croton yellow vein mosaic virus and its cognate betasatellite in germplasm of different Crambe spp in India. Virus Res 243:60–64 Leppik EE (1973) Diseases of crambe. Plant Dis Rptr 57:704–708

Crassocephalum crepidioides (Thickhead) Family: Asteraceae

Ageratum enation virus Taxonomic position Genus: Begomovirus

Leafy vegetable

(AEV)

Family: Geminiviridae

AEV infection in plants of Crassocephalum crepidioides was reported from India (Kumar et al. 2011). The virus-infected thickhead plants exhibit yellow vein symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of AEV, refer to Ageratum spp.

Crassocephalum crepidioides (Thickhead)

661

Crassocephalum yellow vein virus Taxonomic position Genus: Begomovirus

(CraYVV)

Family: Geminiviridae

Geographical distribution CraYVV infection in plants of Crassocephalum crepidioides was reported from China (Chen et al. 2008; Dong et al. 2008). Symptoms and host(s) The virus-infected thickhead plants showed yellow vein symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm, with a single-coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2745 nt (EF165536 = NC_008794) (Briddon 2001; Dong et al. 2008; Brown et al. 2015; Zerbini et al. 2017).

Emilia yellow vein virus Taxonomic position Genus: Begomovirus

(EmYVV)

Family: Geminiviridae

EmYVV infection in plants of Crassocephalum crepidioides was reported from China (Yang et al. 2012). The virus-infected thickhead plants show foliar yellow vein symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of EmYVV, refer to Emilia sonchifolia.

Tobacco leaf curl Yunnan virus Taxonomic position Genus: Begomovirus

(TbLCYnV)

Family: Geminiviridae

TbLCYnV infection in plants of Crassocephalum crepidioides was reported from China (Chen et al. 2008). The virus-infected thickhead plants exhibit leaf curling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of TbLCYnV, refer to Nicotiana tabacum.

References Briddon RW (2001) Begomovirus. Geminiviridae. The Springer index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55

C

662

Crataegus spp. (Hawthorn)

Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Chen J-L, Li F, Li Y, Yi S-X, Guo J, Chen HR (2008) Molecular identification of Geminiviruses inducing vein yellowing in Crassocephalum crepidioides. J Yunnan Agric Univ 23:29–32 Dong JH, Zhang ZK, Ding M, Fang Q, Zhou H (2008) Molecular characterization of a distinct Begomovirus infecting Crassocephalum crepidioides in China. J Phytopathol 156:193–195 Kumar Y, Hallan V, Zaidi AA (2011) First report of Ageratum enation virus infecting Crassocephalum crepidioides (Benth) S. Moore and Ageratum conyzoides L. in India. J Gen Plant Pathol 77:214–216 Yang CX, Luo JS, Zheng LM, Wu ZJ, Xie LH (2012) First report of the occurrence of Emilia yellow vein virus in Crassocephalum crepidioides in China. J Plant Pathol 94:S4.87 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Crataegus spp. (Hawthorn) Family: Rosaceae

Medicinal

Apple chlorotic leaf spot virus Taxonomic position Genus: Trichovirus

(ACLSV)

Family: Betaflexiviridae

ACLSV infection in plants of Crataegus pinnatifida was reported from the UK, China and Greece (Sweet 1980; Katsiani et al. 2014; Wang and Dai 2015; Guo et al. 2016). No vector is reported for this virus. The virus is mechanically sap-transmissible and also transmissible through grafting. For more details of ACLSV, refer to Malus domestica.

References Guo W, Zheng W, Wang M, Li X, Ma Y, Dai H (2016) Genome sequences of three Apple chlorotic leaf spot virus strains from Hawthorns in China. PLoS One. https://doi.org/10.1371/journal.pone.0161099 Katsiani AT, Maliogka VI, Candresse T, Katis NI (2014) Host-range studies, genetic diversity and evolutionary relationships of ACLSV isolates from ornamental, wild and cultivated Rosaceous species. Plant Pathol 63:63–71 Sweet JB (1980) Hedgerow hawthorn (Crataegus spp.) and blackthorn (Prunus spinosa) as hosts of fruit tree viruses in Britain. Ann Appl Biol 94:83–90 Wang M, Dai H (2015) First report of Apple chlorotic leaf spot virus in Hawthorn in China. Plant Dis 99:164

Crinum spp. (Poison bulb) Family: Amaryllidaceae

Ornamental

Hippeastrum mosaic virus Taxonomic position Genus: Potyvirus

(HiMV)

Family: Potyviridae

Crocosmia spp.

663

HiMV was reported in plants of Crinum spp. in the UK (Brunt 1972). The virus-infected poison bulb plants exhibit symptoms of chlorotic streaking in leaves and flower stems. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of HiMV, refer to Hippeastrum spp.

Nerine yellow stripe virus Taxonomic position Genus: Potyvirus

(NeYSV)

Family: Potyviridae

NeYSV infection in plants of Crinum asiaticum was reported from India (Kumar et al. 2015). The virusinfected poison bulb plants exhibit severe mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of NeYSV, refer to Nerine spp.

Tomato zonate spot virus

(TZSV)

Taxonomic position TZSV is a tentative member of the genus Orthotospovirus and family Tospoviridae TZSV infection in plants of Crinum asiaticum was reported from China (Wu et al. 2015). The virus-infected iris plants exhibit symptoms of foliar ringspots and chlorotic spots. The virus is transmitted by thrips vectors in a persistent propagative manner. For more details of TZSV, refer to Solanum lycopersicum.

References Brunt AA (1972) Rep. Glasshouse Crops Res. Inst. For 1972, 103 Kumar S, Raj R, Kaur C, Raj SK (2015) Association of Nerine yellow stripe virus with mosaic disease of Crinum asiaticum ornamental plant in India. Plant Dis 99:1655 Wu K, Zheng KY, Zhang ZK, McBeath JH, Dong JH (2015) First report of Crinum asiaticum as a natural host of Tomato zonate spot virus in China. J Plant Pathol 97:S76

Crocosmia spp. Family: Iridaceae

Ornamental

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

C

664

Crocus spp. (Saffron)

BYMV infection in plants of Crocosmia spp. was reported from the Netherlands (Derks et al. 1980). The virus-infected crocosmia plants exhibit typical foliar mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Cucumber mosaic virus

(CMV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV infection in plants of Crocosmia spp. was reported from Lithuania (Samuitiene and Navalinskiene 2008). The virus-infected crocosmia plants exhibit symptoms of stem distortion, poorly developed flowers, and light green streaks on leaves. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

References Derks AFLM, Vink-van den Abeele JL, Muller PJ (1980) Bean yellow mosaic virus in some iridaceous plants. Acta Hortic 110:31–38 Samuitiene M, Navalinskiene M (2008) Occurrence of Cucumber mosaic cucumovirus on ornamental plants in Lithuania. Zemdirbyste-Agriculture 95:135–143

Crocus spp. (Saffron) Family: Iridaceae

Ornamental

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

ArMV infection in plants ofIridaceae Crocus spp. was reported from England and Lithuania (Loebenstein et al. 1995; Samuitiene et al. 2008). The petals of blue-colored cultivars of saffron plants infected with ArMV show white streaks extending toward the edge of the petals. The petals are crinkled and wavy, but the leaves remain symptomless. This virus is transmitted by nematode vectors (Xiphinema and Dorylamidae spp.) in a non-persistent manner. The virus is also transmitted by mechanical sap inoculation, grafting, and seed. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

Crocus spp. (Saffron)

665

BYMV infection in plants of Crocus spp. was reported from Italy, the Netherlands, and Japan (Russo et al. 1979; Derks and Vink-Van den Abeele 1982; Kaneshige et al. 1991; Loebenstein et al. 1995; Grilli Caiola and Faoro 2011). The virus-infected saffron plants show symptoms of a faint mosaic on the leaves after flowering; later in the season, leaf symptoms disappear. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Crocus spp. was reported from Lithuania and Netherlands (Derks 1995; Chen et al. 2001; Navalinskiene and Samuitiene 2001; Chen 2003; Samuitiene and Navalinskiene 2008). The virus-infected saffron plants exhibit foliar mosaic and streaking symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sapinoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Iris mild mosaic virus Taxonomic position Genus: Potyvirus

(IMMV)

Family: Potyviridae

IMMV-infected in plants of Crocus spp. plants were reported from Lithuania (Navalinskiene and Samiutiene 2001). The virus-infected saffron plants show narrowed, twisted leaves with light green streaking and occasional necrotic spotting and streaking. Petals showed crinkling and color break. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of IMMV, refer to Iris spp.

Iris severe mosaic virus Taxonomic position Genus: Potyvirus

(ISMV)

Family: Potyviridae

ISMV infection in plants of Crocus tomasinianus was reported from the Netherlands (Brunt and Phillips 1979). The virus-infected saffron plants exhibit a mosaic of light green and small stripes on the leaves. The flowers of blue-colored cultivars show dark blue stripes. The aphids Myzus persicae and Macrosiphum euphorbiae transmit the virus in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of ISMV, refer to Iris spp.

Narcissus mosaic virus Taxonomic position Genus: Potexvirus

(NMV)

Family: Alphaflexiviridae

C

666

Crocus spp. (Saffron)

NMV was reported in plants of Crocus spp. from Netherlands by Miglino et al. (2005). The virusinfected saffron plants exhibit symptoms of stunting, yellowing, necrosis, and flower color breaking. The virus is mechanically sap-transmissible. For more details of NMV, refer to Narcissus spp.

Saffron latent virus Taxonomic position Genus: Potyvirus

(SaLV)

Family: Potyviridae

Geographical distribution SaLV infection in plants of Crocus sativus was reported Iran (Parizad et al. 2017) Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is transmitted by mechanical sap inoculation. The virus is not transmitted by contact between plants; not transmitted by seed; not transmitted by pollen. Virion properties and genome The virions are non-enveloped, flexuous filaments 680–900 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 9693 nt (KY562565 = NC_036802) excluding the poly-A tail (Revers and Garcia 2015; Parizad et al. 2017; Wylie et al. 2017).

Tobacco necrosis virus

(TNV)

Taxonomic position TNV is a tentative member of the genera Necrovirus and and family Tombusviridae. TNV was reported occasionally in Crocus spp. plants in the Netherlands (Miglino et al. 2005) and in Lithuania (Navalinskiene and Samuitiene 2006). The virus-infected saffron plants exhibit symptoms of leaves that were smaller than in healthy plants, showing distortion, indented margins, chlorotic streaks, and necrotic spots. The virus is transmitted by the zoospores of the fungus Olpidium brassicae, and the virus is also mechanically sap-transmissible. For more details of TNV, refer to Nicotiana tabacum.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in plants of Crocus spp. was reported from Lithuania (Navalinskiene and Samuitiene 2001). The virus-infected saffron plants exhibit yellowish-green to white spots and stripes on the leaves which are notched. The flowers may show dark spots (Asjes 1974). The virus is transmitted by nematode vectors and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Crocus spp. (Saffron)

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

667

(TRSV)

Family: Secoviridae

TRSV has rarely been reported in plants of Crocus spp. in the Netherlands (Derks 1995). The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Crocus sativus was reported from China and New Zealand (Chen and Chen 2000; Ochoa Corona et al. 2007). The virus-infected saffron plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors, Myzus persicae, Brevicoryne brassicae, and other aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TuMV, refer to Brassica rapa.

References Asjes CJ (1974) Soil-borne viral diseases in ornamental bulbous crops and their control in the Netherlands. Agric Environ 1:303–315 Brunt AA, Phillips S (1979) Viruses of bulb crops, Crocus tomasinianus and Narcissus spp. Annual Report of the Glasshouse crops Horticultural Research Institute 1979, pp 130–131 Chen YK (2003) Occurence of cucumber mosaic virus in ornamental plants and perspectives of transgenic control. Ph.D. Thesis, Wageningen University, The Netherlands, p 144 Chen J, Chen JS (2000) Occurrence and control of mosaic disease (Turnip mosaic virus) in saffron (Crocus sativus). Zhejiang Nongye Kexue 3:132–135 Chen YK, Derks AF, Langeveld S, Goldbach R, Prins M (2001) High sequence conservation among cucumber mosaic virus isolates from lily. Arch Virol 146:1631–1636 Derks AFLM (1995) Crocus spp. Chapter 21. In: Loebenstein G, Lawson RH, Brunt AA (eds) Virus and virus-like diseases of bulb and flower crops. Wiley, Chichester Derks AFLM, Vink-Van den Abeele JL. (1982). Jversl. Lab. Bloembollen Onderz., Lisse 1981:51–53 Grilli Caiola M, Faoro F (2011) Latent virus infections in Crocus sativus and Crocus cartwrightianus. Phytopathol Mediterr 50:175–182 Kaneshige H, Maeda T, Inouye N (1991) Host range and properties of Bean yellow mosaic virus (BYMV) infecting crocus, and serological relationships among three strains of BYMV. Nogaku Kenkyu 62:225–240 Loebenstein G, Lawson RH, Brunt AA (1995) Virus and virus-like diseases of bulb and flowering crops. Wiley, UK, p 543 Miglino R, Jodlowska A, van Schadewijk AR (2005) First report of Narcissus mosaic virus infecting Crocus spp. cultivars in the Netherlands. Plant Dis 89:342 Navalinskiene M, Samuitiene M (2001) Viruses affecting some bulb and corn flower crops. Biologia 4:40–42 Navalinskiene M, Samuitiene M (2006) Dekoratyvinių augalų virusin_es ligos ir jų suk_el_ejai Lietuvoje. Lutut_e, Kaunas 254 p Ochoa Corona FM, Lebas BSM, Elliott DR, Tang JZ, Alexander BJR (2007) New host records and new host family range for Turnip mosaic virus in New Zealand. Austral Plant Dis Notes 2:127–130 Parizad S, Dizadji A, Koobi Habibi M, Winter S, Kalantari S, Garcia-Arenal F, Ayllon MA (2017) Prevalence of Saffron latent virus (SaLV), a new Potyvirus species, in saffron fields of Iran. J Plant Pathol 99:802 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199

C

668

Crossandra infundibuliformis (Firecracker flower)

Russo M, Martelli GP, Cresti M, Ciampolini F (1979) Bean yellow mosaic virus in saffron. Phytopathol Mediterr 18:189–191 Samuitiene M, Navalinskiene M (2008) Occurrence of Cucumber mosaic cucumovirus on ornamental plants in Lithuania. Zemdirbyste-Agriculture 95:135–143 Samuitiene M, Navalinskiene M, Jackeviciene E (2008) Arabis mosaic virus on ornamental plants. Biologia 54:264–268 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Crossandra infundibuliformis (Firecracker flower) Family: Acanthaceae

Ornamental

Alternanthera mosaic virus Taxonomic position Genus: Potexvirus

(AltMV)

Family: Alphaflexiviridae

AltMV infection in plants of Crossandra infundibuliformis was reported from Florida and other locations in the USA (Baker et al. 2006). The virus-infected firecracker flower plants exhibit mosaic symptoms. No vector is reported for this virus. The virus is readily sap-transmissible and easily spread by plant contact, vegetative propagation, and contaminated tools and hands. For more details of AltMV, refer to Alternanthera spp.

References Baker CA, Breman L, Jones L (2006) Alternanthera mosaic virus found in Scutellaria, Crossandra, and Portulaca in Florida. Plant Dis 90:833

Crotalaria spp. (Sunn hemp, Showy rattlebox) Family: Fabaceae

Weed host

Bean common mosaic necrosis virus Taxonomic position Genus: Potyvirus

(BCMNV)

Family: Potyviridae

BCMNV was reported in plants of Crotalaria incana from Uganda by Sengooba et al. (1997). The virus-infected sunn hemp plants exhibit mild mosaic symptoms. The virus is transmitted by an aphid vector, Aphis fabae. in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BCMNV, refer to Phaseolus vulgaris.

Crotalaria spp. (Sunn hemp, Showy rattlebox)

Bean common mosaic virus Taxonomic position Genus: Potyvirus

669

(BCMV)

Family: Potyviridae

BCMV infection in plants of Crotalaria striata was reported from India (Sarkar and Kulsheshtra 1978). The virus-infected sunn hemp plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BCMV, refer to Phaseolus vulgaris.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Crotalaria spectabilis was reported from Japan (Komuro and Iwaki 1968). The virus-infected sunn hemp plants exhibit yellow mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Cowpea aphid-borne mosaic virus Taxonomic position Genus: Potyvirus

(CABMV)

Family: Potyviridae

CABMV infection in plants of Crotalaria juncea was reported from Nigeria (Ladipo et al. 2005; Gonzalez Segnana et al. 2013). The virus-infected sunn hemp plants exhibit mosaic, distortion, and puckering symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CABMV, refer to Vigna unguiculata.

Cowpea mosaic virus Taxonomic position Genus: Comovirus

(CPMV)

Family: Secoviridae

CPMV infection in plants of Crotalaria juncea was reported from Nigeria (Ladipo 1988). The virusinfected sunn hemp plants exhibit mosaic, distortion, and puckering symptoms. The virus is transmitted by beetle vectors and also by mechanical sap-inoculation. For more details of CPMV, refer to Vigna unguiculata.

Cowpea severe mosaic virus Taxonomic position Genus: Comovirus

(CPSMV)

Family: Secoviridae

C

670

Crotalaria spp. (Sunn hemp, Showy rattlebox)

CPSMV infection in plants of Crotalaria spp. was reported from Brazil (Lin et al. 1982; Lima et al. 2005). The virus-infected sunn hemp plants exhibit chlorotic mottling and leaf distortion or with chlorotic spot symptoms. The virus is transmitted by the beetle vectors and also through mechanical sap-inoculation. For more details of CPSMV, refer to Vigna unguiculata.

Sunn hemp leaf distortion virus Taxonomic position Genus: Begomovirus

(SHLDV)

Family: Geminiviridae

Geographical distribution SHLDV infection in plants of Crotalaria juncea was reported from Barrackpore (India) (Das et al. unpublished - FJ455449). Symptoms and host(s) The virus-infected sunn hemp plants exhibit leaf distortion symptoms. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm, with a single-coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2774 nt (FJ455449 = NC_013019) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Sunn-hemp mosaic virus Taxonomic position Genus: Tobamovirus

(SHMV)

Family: Virgaviridae

Geographical distribution SHMV was first reported in plants of Crotalaria juncea from India by Capoor and Varma (1948). The virus spreads in India and the USA (Silver et al. 1996). Symptoms and host(s) The virus-infected sunn hemp plants exhibit systemic mosaic, mottling, and stunting symptoms. Transmission The virus is not transmitted by any insect vector. The virus is transmissible by mechanical sapinoculation, transmissible by contact between plants, and probably transmissible by seed but not transmissible by pollen. Virion properties and genome The virions are non-enveloped, rigid helical rods with a helical symmetry, about 17 nm in diameter, and 300 nm in length. The genome is a monopartite, linear, and positive-sense ssRNA of 6.3–6.5 kb. Partial genome sequences are available (U47034; J02413). The 30 -terminus has a tRNA-like structure.

Crotalaria spp. (Sunn hemp, Showy rattlebox)

671

The 50 terminus has a methylated nucleotide cap (m7G50 pppG) (Silver et al. 1996; Zaitlin 2011; Adams et al. 2017).

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Crotalaria spectabilis was reported from Japan (Komuro and Iwaki 1968). The virus-infected sunn hemp plants exhibit yellow ringspot and mosaic symptoms. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Crotalaria spp. was reported from India and Australia (Bhat et al. 2002; Prasad et al. 2005; Sharman et al. 2015). The virus-infected sunn hemp plants showed chlorotic/ necrotic spots on young leaves and petioles and bending and necrosis of the shoot. The virus is transmitted by the thrips vectors, the virus present in/on pollen, and entering in to the host through the injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato severe rugose virus Taxonomic position Genus: Begomovirus

(ToSRV)

Family: Geminiviridae

ToRSV infection in plants of Crotalaria spp. was reported from Brazil (Barreto et al. 2013). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ToSRV, refer to Solanum lycopersicum.

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV infection in plants of Crotalaria spectabilis was reported from Florida, USA (Adlerz 1969). The virus-infected sunn hemp plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of WMV, refer to Citrullus lanatus.

C

672

Croton bonplandianum (Ban tulsi)

References Adams MJ, Adkins S, Bragard C, Gilmer D, Li D, MacFarlane SA, Wong S-M, Melcher U, Ratti C, Ryu KH, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Virgaviridae. J Gen Virol 98:1999–2000 Adlerz WC (1969) Distribution of watermelon mosaic viruses 1 and 2 in Florida. Fla Agric Exp Station J 3408:161–165 Barreto SS, Hallwass M, Aquino OM, Inoue-Nagata AK (2013) A study of weeds as potential inoculum sources for a tomato-infecting begomovirus in central Brazil. Phytopathology 103(5):436–444 Bhat AI, Jain RK, Chaudhury V, Krishnareddy M, Ramaiah K, Chattannavar SN, Varma A (2002) Sequence conservation in the coat protein gene of Tobacco streak virus isolates causing necrosis disease in cotton, mung bean, sunflower, and sunn-hemp in India. Indian J Biotecnol 1:350–356 Briddon RW (2001) Begomovirus. Geminiviridae. The Springer index of Viruses. Springer, New York, p 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Capoor SP, Varma PM (1948) Enation mosaic of Dolichos lablab Linn, a new virus disease. Curr Sci 17:57 Gonzalez Segnana LR, Farina AE, Gonzalez DD, Mello APOA, Rezende JAM, Kitajima EW (2013) Alternate hosts of Cowpea aphid-borne mosaic virus (CABMV) in sesame (Sesamum indicum) crops grown in Paraguay. Trop Plant Pathol 38:539–542 Komuro Y, Iwaki M (1968) Bean yellow mosaic virus and tobacco ringspot virus isolated from Crotalaria (Crotalaria spectabilis). Ann Phytopathol Soc Jpn 34:7–15 Ladipo JL (1988) Viruses associated with a mosaic disease of Crotalaria juncea in Nigeria. J Phytopathol 121:8–18 Ladipo JL, Lommel SA, Barnett OW (2005) Identification and characterization of cowpea aphid-borne mosaic virus as the second virus from mixed-infected Crotalaria juncea plants in Nigeria. Z Pflanzenkrankh Pfalnzenschutz 112:222–228 Lima JA, Nacimento AKQ, Silva GS, Camarco RFEA, Goncalves MFB (2005) Crotalaria paulinea, a new natural host of Cowpea severe mosaic virus. Fitopatol Bras 30:429–433 Lin MT, Anjos JRN, Rios GP (1982) Cowpea severe mosaic virus in five legumes in central Brazil. Plant Dis 66:67–70 Prasad MSL, Madhavi J, Prasada Rao RDVJ, Singh H, Marimutu T (2005) Incidence of Tobacco streak virus on Sunn hemp (Crotalaria juncea L.). Indian J Plant Protect 33:297 Sarkar KR, Kulsheshtra K (1978) Crotalaria striata DC., a new natural host of Bean common mosaic virus. Curr Sci 47:241 Sengooba TN, Spence NJ, Walkey DGA, Allen DJ, Femi Lana A (1997) The occurrence of Bean common mosaic necrosis virus in wild and forage legumes in Uganda. Plant Pathol 46:95–103 Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct Tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207 Silver S, Quan S, Deom CM (1996) Completion of the nucleotide sequence of sunn-hemp mosaic virus: a tobamovirus pathogenic to legumes. Virus Genes 13(1):83–85 Zaitlin M (2011) Tobamovirus. In: Tidona C, Darai G (eds) The Springer index of viruses. Springer, New York. https://doi. org/10.1007/978-0-387-95919-1 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Croton bonplandianum (Ban tulsi) Synonyms Croton sparsiflorus Family: Euphorbiaceae

Ageratum enation virus Taxonomic position Genus: Begomovirus

Weed host

(AEV)

Family: Geminiviridae

Croton bonplandianum (Ban tulsi)

673

AEV infection in plants of Croton bonplandianum was reported from India (Khan et al. 2014). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of AEV, refer to Ageratum spp.

Croton yellow vein mosaic virus Taxonomic position Genus: Begomovirus

(CroYVMV)

Family: Geminiviridae

Geographical distribution CroYVMV first reported in plants of Croton bonplandianum from India by Varma (1963) and subsequently by Mandal (1989) and Mandal and Muniyappa (1991a, b). Symptoms and host(s) The virus-infected ban tulsi plants exhibit typical symptoms of vein thickening, severe inward curling and a reduction in leaf size, and stunting. In certain varieties pink or yellow veins with foliar malformation were also observed. In addition to C. bonplandianum, CroYVMV is known to cause natural infection in several weeds and economic plant species, e.g., Jatropha spp., Acalypha spp., Cyamopsis spp., Crotalaria juncea, okra, papaya, radish, rapeseed-mustard, and tomato. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is graft-transmissible (Mandal and Muniyappa 1991b). CroYVMV has a large experimental host range. It infects as many as 35 plant species including vegetables, tobacco, ornamentals, and weeds of 11 families. Virion properties and genome In leaf-dip and purified preparations of CroYVMV-infected C. bonplandianum, geminate virions of 18–30 nm were observed (Mandal and Muniyappa 1991b). The genome consists of a single molecule of circular single-stranded DNA. The analysis of complete genome sequences of four croton isolates from northern, eastern, and southern India showed the association of a single species of DNA-A contains 2757 nt (AJ507777 = NC_004300, EU727086, JN831446, JN817516, JN817517). Croton yellow vein mosaic betasatellite DNA molecule is associated with CroYVMV, and consists of 1346 nt (AM410551; JQ354987, JN831447, JN831448) (Briddon 2001; Mandal et al. 2001; Hussain et al. 2011; Pramesh et al. 2013; Zhou 2013; Brown et al. 2015; Zerbini et al. 2017).

Papaya leaf curl virus Taxonomic position Genus: Begomovirus

(PaLCuV)

Family: Geminiviridae

PaLCuV infection in plants of Croton bonplandianum was reported from India (Chowda Reddy et al. unpublished - AY738092; Jaidi et al. unpublished - KM525657). The virus-infected ban tulsi plants exhibit yellow vein and leaf curl symptoms. The virus is transmitted by the whitefly vector, Bemisia

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Cryptocoryne spp. (Water trumpet)

tabaci, in a circulative non-propagative manner. The virus is not mechanically sap-transmissible. For more details of PaLCuV, refer to Carica papaya.

References Briddon RW (2001) Begomovirus. Geminiviridae. The Springer index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55. Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Hussain K, Hussain M, Mansoor S, Briddon RW (2011) Complete nucleotide sequence of a Begomovirus and associated betasatellite infecting croton (Croton bonplandianus) in Pakistan. Arch Virol 156:1101–1105 Khan MS, Tiwari AK, Raj SK, Srivastava A, Hye Ji S, Chun SC (2014) Molecular epidemiology of begomoviruses occurring on some vegetables, grain legume and weed species in the Terai belt of north India. J Plant Dis Protect 121:53–57 Mandal B (1989) Studies on Croton yellow vein mosaic virus. MSc. Thesis. University of Agricultural Sciences, Bangalore, India. 132 pp Mandal B, Muniyappa V (1991a) Isolation and detection of Croton yellow vein mosaic virus, a whitefly transmitted Geminivirus. Fitopatol Bras 16:177–180 Mandal B, Muniyappa V (1991b) Transmission of Croton yellow vein mosaic virus by Bemisia tabaci: virus-vector relationship. Fitopatol bras 16:260–262 Mandal B, Pappu HR, Muniyappa V (2001) Characterization of Croton yellow vein mosaic virus, family Geminiviridae. Phytopathology 91(6):S57 Pramesh D, Mandal B, Phaneendra C, Muniyappa V (2013) Host range and genetic diversity of Croton yellow vein mosaic virus, a weed-infecting monopartite Begomovirus causing leaf curl disease in tomato. Arch Virol 158:531–542 Varma PM (1963) Transmission of plant viruses by whiteflies. Natl Inst Sci India Bull 24:11–33 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zhou X (2013) Advances in understanding Begomovirus satellites. Annu Rev Phytopathol 51:357–381

Cryptocoryne spp. (Water trumpet) Family: Araceae

Dasheen mosaic virus Taxonomic position Genus: Potyvirus

Ornamental

(DsMV)

Family: Potyviridae

DsMV infection in plants of Cryptocoryne spp. was reported from Hawaii (Nelson 2008). The virusinfected water trumpet plants show symptoms such as systemic mosaic, leaf malformation, epinasty, veinclearing, local and systemic necrosis. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of DsMV, refer to Colocasia esculenta.

References Nelson SC (2008) Dasheen mosaic of edible and ornamental aroids. Plant Disease CTAHR. Available at: http://www. ctahr.hawaii.edu/oc/freepubs/pdf/PD-44.pdf

Cucumis anguria (Gherkin)

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Cryptotaenia japonica (Japanese hornwort) Family: Apiaceae

Leafy vegetable

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

C

Family: Bromoviridae

CMV infection in plants of Cryptotaenia japonica was reported from Japan (Komuro and Asuyama 1955; Fujita et al. 1999). The virus-infected Japanese hornwort plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Konjac mosaic virus

(KoMV)

Synonyms Japanese hornwort mosaic virus (JHMV) Taxonomic position Genus: Potyvirus

Family: Potyviridae

JHMV infection in plants of Cryptotaenia japonica was reported from Japan (Okuno et al. 2003). The virus-infected Japanese hornwort plants exhibit mosaic symptoms. The virus is transmitted by the aphid vector, Myzus persicae, in a non-persistent manner, and also by mechanical sap-inoculation. The host range for this virus was narrow and it could systemically infected only plants belonging to Umbelliferae (Apiaceae). For more details of KoMV, refer to Amorphophallus paeoniifolius.

References Fujita T, Saiga T, Yamagata T, Harada Y (1999) The virus belongs to Potyviridae detected from Cryptotaenia japonica plants with mosaic symptom. Ann Phytopathol Soc Jpn 65:661 Komuro Y, Asuyama H (1955) Studies on cucumber mosaic virus. II. The isolation of the virus from various plants showing mosaic in the vicinity of Tokyo, Japan. Ann Phytopathol Soc Jpn 20:77–82 Okuno K, Hama T, Takeshita M, Furuya N, Takanami Y (2003) New Potyvirus isolated from Cryptotaenia japonica. J Gen Plant Pathol 69:138–142

Cucumis anguria (Gherkin) Family: Cucurbitaceae

Vegetable

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

(CGMMV)

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Cucumis anguria (Gherkin)

CGMMV infection in plants of Cucumis anguria was reported from India (Reshmi et al. 2005a). The virus-infected gherkin plants exhibit symptoms of yellowing, stunted growth, blistering, mosaic mottling, chlorotic spots, necrotic lesions, and leaf cupping. The virus is transmissible by mechanical sap-inoculation, and also by contact between plants. Transmission through contaminated irrigation water or nutrient solutions has also been reported. The virus is also pollen transmissible. For more details of CGMMV, refer to Cucumis sativus.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Cucumis anguria was reported from southern Karnataka (India) (Kavyashri 2014; Kavyashri and Nagaraju 2014). The virus is transmitted by aphid vector, Aphis gossypii, in a nonpersistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Papaya ringspot virus Taxonomic position Genus: Potyvirus

(PRSV)

Family: Potyviridae

PRSV-P pathotype infection in plants of Cucumis anguria was reported from Bangalore (India) (Reshmi et al. 2005b). The aphid vector, Myzus persicae, is an efficient vector for this virus and the virus is transmissible by mechanical sap-inoculation. For more details of PRSV, refer to Carica papaya.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Cucumis anguria was reported from India (Krishnareddy et al. 2003; Sarovar et al. 2010). The virus-infected gherkin plants exhibit necrosis on leaves, stem, flower buds, and shoot tip resulting in death of veins and dieback. The virus is transmitted by thrips vectors, the virus present in/on pollen, and entering into the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible. The virus is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

Cucumis melo (Muskmelon or Cantaloupe)

677

WMV infection in plants of Cucumis anguria was reported from Venezuela (Romay et al. 2013). The virus-infected gherkin plants exhibit systemic mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of WMV, refer to Citrullus lanatus.

Zucchini yellow mosaic virus Taxonomic position Genus: Potyvirus

(ZYMV)

Family: Potyviridae

ZYMV infection in plants of Cucumis anguria was reported from southern India (Andhra Pradesh) and the Czech Republic (Svoboda and Polak 2002; Johnson et al. 2013). The virus-infected gherkin plants show mosaic, blistering of leaves, and malformed fruits. The virus is transmitted by a wide range of aphid species of which the green peach (Myzus persicae) and melon (Aphis gossypii) aphids are the most important and also by mechanical sap-inoculation. For more details of ZYMV, refer to Cucurbita pepo.

References Johnson AMA, Vidya T, Papaiah S, Srinivasulu M, Mandal B, Sai Gopal DVR (2013) First Report of Zucchini yellow mosaic virus Infecting Gherkin (Cucumis anguira) in India. Indian J Virol 24:289–290 Kavyashri VV (2014) Survey, biological and molecular characterization and management of Cucumber mosaic virus in Gherkin (Cucumis anguria L.). Thesis submitted to the Department of Plant Pathology, UAS, GKVK, Bangalore Kavyashri VV, Nagaraju N (2014) Molecular survey for incidence of Cucumber mosaic virus in gherkin (Cucumis anguria L.) and its transmission. Mysore J Agric Sci 48(3):381–386 ref.16 Krishnareddy M, Devaraj LR, Salil J, Samuel DK (2003) Outbreak of Tobacco streak virus causing necrosis of cucumber (Cucumis sativus) and Gherkin (Cucumis anguria) in India. Plant Dis 87:1264 Reshmi CM, Lakshmi Narayana Reddy CN, Praveen HM, Nagaraju (2005a) Natural occurrence of Cucumber green mottle mosaic virus (CGMMV) in Gherkin (Cucumis anguria L.). Environ Ecol 23(SPL 4):781–784 Reshmi CM, Krishnareddy M, Praveen HM, Nagaraju (2005b) Occurrence of Papaya ring spot virus (PRSV-P) on gherkin (Cucumis anguria L.). Environ Ecol 23S:785–788 Romay G, Lecoq H, Desbiez C (2013) First report of watermelon mosaic virus naturally infecting Cucumis anguria. Plant Dis 97:1515 Sarovar B, Prasad YS, Sai Gopal DVR (2010) Detection of Tobacco streak virus by immunocapture-reverse transcriptasepolymerase chain reaction and molecular variability analysis of a part of RNA3 of sunflower, gherkin, and pumpkin from Andhra Pradesh, India. Science Asia 36:194–198 Svoboda J, Polak J (2002) Distribution, variability and overwintering of Zucchini yellow mosaic virus in the Czech Republic. Plant Protect Sci 38:125–130

Cucumis melo (Muskmelon or Cantaloupe) Family: Cucurbitaceae

Beet pseudoyellows virus Synonyms Cucumber yellows virus

Edible fruit

(BPYV)

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Taxonomic position Genus: Crinivirus

Cucumis melo (Muskmelon or Cantaloupe)

Family: Closteroviridae

BPYV infection in plants of Cucumis melo was reported from France, Spain, Bulgaria, Greece, and Italy (Tomassoli et al. 2003; Berdiales et al. 1999; Boubourakas et al. 2006). The virus-infected muskmelon plants first exhibit symptoms on intermediate, and lower leaves with angular chlorotic spots that expand and coalesce inducing a generalized yellowing of interveinal tissue, while the veins remain green. Old yellow leaves were thick and brittle producing a characteristic snap when crushed, while young leaves and vegetative tips often appeared normal. In symptomatic plants, fruits stopped growing, and ripening and yields were seriously compromised. The virus is transmitted by the greenhouse whitefly Trialeurodes vaporariorum in a semi-persistent manner (Yamashita et al. 1979; Wisler et al. 1998; Caciagli 2001). The virus is not transmissible by mechanical sap-inoculation, and also not by contact between plants. For more details of BPYV, refer to Beta vulgaris.

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

CGMMV infection in plants of Cucumis melo was reported from China, Japan, Israel, Pakistan, New Delhi (India), and California (USA) (Raychaudhuri and Varma 1978; Yoshida et al. 1980b; Antignus et al. 1990; Zhang et al. 2009; Ali et al. 2014; Tian et al. 2014). The virus-infected muskmelon plants exhibit symptoms of light yellow-green spots and vein-clearing on young leaves to chlorotic mottling, plant stunting, and necrosis at later stages of growth. There is no known vector for this virus. The virus is transmissible by mechanical sap-inoculation. Vani and Varma (1993) have reported a positive role of irrigation water from Jamuna River (New Delhi, India) in the spread of CGMMV under natural conditions. The virus is seed and pollen transmissible in Cucumis melo plants (Wu et al. 2011). For more details of CGMMV, refer to Cucumis sativus.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Cucumis melo was reported from China, Bosnia, California, Spain, Iran, Japan, and Herzegovina (Yoshida et al. 1980b; Tobias et al. 1982; Sharma et al. 1984; Rivera et al. 1991; Grafton-Cardwell et al. 1996; Luis-Arteaga et al. 1998; Ozaslan et al. 2006; Malik et al. 2010; Lima et al. 2012; Trkulja et al. 2013; Romay et al. 2014; Rasoulpour et al. 2016; Wang et al. 2017). The virus-infected muskmelon plants exhibit symptoms of mosaic, chlorotic mottling, and vein-banding as well as blistering and leaf malformation. The virus is transmitted by a number of aphids in a non-persistent manner (Garzo et al. 2004). The virus is also mechanically sap-transmissible to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Cucumis melo (Muskmelon or Cantaloupe)

Cucumber vein yellowing virus Taxonomic position Genus: Ipomovirus

679

(CVYV)

Family: Potyviridae

CVYVoccurs very commonly on Cucumis melo in the Mediterranean basin, Iran, France, Portugal, and Tunisia (Yilmaz et al. 1989; Louro et al. 2004; Bananej et al. 2006; Lecoq et al. 2007; Yakoubi et al. 2007). The virus-infected muskmelon plants exhibit symptoms of yellowing of the leaf vein area, becoming systemic, with chlorosis of the youngest leaves. In extreme infections, plant stunting and sudden death of plants have been described. Affected fruits show chlorotic spots on their skin and/or internal necrosis. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CVYV, refer to Cucumis sativus.

Cucumis melo alphaendornavirus Taxonomic position Genus: Alphaendornavirus

(CmEV)

Family: Endornaviridae

Geographical distribution CmEV infection in plants of Cucumis melo was reported from the USA (Sabanadzovic et al. 2016). Symptoms and host(s) The virus-infected muskmelon plants do not exhibit any external symptoms. Transmission No vector has been identified and virus is not mechanically sap-transmissible. The virus is transmissible through seed via both ova and pollen. Virion properties and genome The virus does not have a gene for a capsid protein; therefore no virion is formed. The genome is a double-stranded RNA consisting of 15,078 bp and contains a single 4939 codons-long ORF that terminates with a stretch of 10 cytosine residues (KT727022).

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(CABYV)

Family: Luteoviridae

Geographical distribution CABYV was first reported from France (Lecoq et al. 1992). The virus infection in Cucumis melo plants was also noticed in China, North America, Taiwan, Algeria, Greece, Turkey, Lebanon, Spain, Italy, Tunisia, Egypt, Montenegro, and Korea (Lemaire et al. 1993; Lecoq et al. 1994; Abou-Jawdah et al. 1997; Deng et al. 1997; Lecoq 1999; Juarez et al. 2004; Hattab et al. 2005; Kassem et al. 2007; Tomassoli and Meneghini 2007; Yardimci and Ozgonen 2007; Xiang et al. 2008a; Hattab et al. 2009; Knierim et al. 2010; Omar and Bagdady 2012; Kassem et al. 2013; Choi et al. 2015; Choi and Choi 2016; Zindovic et al. 2017).

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Cucumis melo (Muskmelon or Cantaloupe)

Symptoms and host(s) The virus-infected muskmelon plants exhibit yellowing of the older leaves, and leaves are thickened and brittle. A wide range of symptom intensity exists depending on the cultivars, varying from yellowing and limited to a few older leaves to a complete discoloration of plants (Lecoq et al. 1992). The virus infection does not affect fruit quality but rather induces flower abortions and reduces the number of fruit per plant. Natural infection of CABYV has been detected in nine different cucurbit species: cushaw (Cucurbita moschata), squash (Cucurbita pepo), calabash gourd (Lagenaria siceraria), cucumber (Cucumis sativus), suakwa vegetable sponge (Luffa cylindrica), watermelon (Citrullus lanatus), muskmelon (Cucumis melo), wax gourd (Benincasa hispida), and bitter gourd (Momordica charantia) in China (Xiang et al. 2008a). Ecballium elaterium is a potential virus reservoir, and melon and squash are the other natural hosts of CABYV in Spain (Kassem et al. 2013). Transmission The virus is transmitted by aphid vectors, Myzus persicae, M. euphorbiae, and Aphis gossypii in a circulative, non-propagative manner (Lecoq et al. 1992; Kassem et al. 2013; Choi and Choi 2016). The virus is not mechanically sap-transmissible. Virion properties and genome The virions are 25–30 nm in diameter, hexagonal in outline, and have no envelope. The genome is a single positive-sense, single-stranded RNA of 5669 nt (NC_003688) (Lecoq et al. 1992). The virus genome codes for six proteins, including two structural proteins, the coat protein of 23 kDa, and a readthrough protein of 76 kDa involved in aphid transmission (Guilley et al. 1994; Xiang et al. 2008b). The 30 terminus has neither a poly (A) tract nor a tRNA-like structure. The 50 terminus has a genome-linked protein (VPg) (of about 17 kDa). The genome contains six open reading frames (ORFs) (Bananej et al. 2009).

Cucurbit chlorotic yellows virus

(CCYV)

Taxonomic position CCYV is a tentative member of the genus Crinivirus and family Closteroviridae Geographical distribution CCYV infection in plants of Cucumis melo was reported from China, Taiwan, Greece, Iran, Japan, and Sudan (Gyoutoku et al. 2009; Huang et al. 2010; Gu et al. 2011; Hamed et al. 2011; Zeng et al. 2011; Bananej et al. 2013; Orfanidou et al. 2014, 2017; Abrahamian and Abou-Jawdah 2014). Symptoms and host(s) The virus-infected muskmelon plants exhibit symptoms of chlorosis, yellowing, and interveinal chlorotic spots on lower leaves. Transmission The virus is transmitted by Bemisia tabaci biotype B and Q whiteflies in a semi-persistent manner (Gyoutoku et al. 2009; Orfanidou et al. 2017). CCYV systemically infected 19 of 21 cucurbitaceous species tested by whitefly transmission; only Cucumis anguira and Cucumis zeyheri were not infected (Okuda et al. 2010). Citrullus lanatus, Cucurbita pepo, and Luffa cylindrica were susceptible to CCYV; however, the infection rates were

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low and symptoms were unclear. In addition to the cucurbitaceous plants, Beta vulgaris, Chenopodium amaranticolor, Chenopodium quinoa, Spinacia oleracea, Lactuca sativa, Datura stramonium, and Nicotiana benthamiana were also systemically infected by CCYV. Virion properties and genome The virions are flexuous filaments of 800–850 nm long and 12 nm wide, and they have a bipartite single-stranded RNA genome. Both genomic RNAs are both required for infectivity, RNA1 of 8607 nt (NC_018173) and RNA2 of 8041 nt (NC_018174), are encapsidated separately in two particles (Okuda et al. 2010; Lin et al. 2012; Tzanetakis et al. 2013).

Cucurbit leaf crumple virus

(CuLCrV)

Synonyms Cucurbit leaf curl virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution CuLCrV infection in plants of Cucumis melo was reported from Arizona, Florida, California and Texas (USA), and Mexico (Kuo et al. 2007; Adkins et al. 2009; Brown et al. 2000; Guzman et al. 2000; Hernandez et al. 2000). Symptoms and host(s) The virus-infected muskmelon plants exhibit leaf curl and/or mottling symptoms. The virus infects most cucurbits including cucumber, muskmelon, squash, pumpkin, and watermelon and has been reported to infect bean (Guzman et al. 2000). Transmission The virus is transmitted in a circulative non-propagative manner by various biotypes of the whitefly, Bemisia tabaci, including the silverleaf whitefly (B. tabaci biotype B). The adult whitefly must feed for a minimum of 30 min on the infected plant and can only transmit the virus after a delay of 6–8 h. Once the whitefly is able to transmit the virus, it can continue to do so for days (Brown et al. 2000). The virus cannot be transmissible mechanically and is unlikely to be transmissible through seed. Virion properties and genome The structure of the virions of CuLCrV has not been investigated. In common with all geminiviruses, the virions of CuLCuV are likely geminate (twinned quasi-icosahedra). CuLCuV is a typical bipartite Begomovirus native to the New World. The genome consists of two circular, single-stranded DNA components. DNA-A contains 2632 nt (AF224760 = NC_002984; AF256200) and DNA-B of 2600 nt (AF224761 = NC_002985; AF327559) (Briddon 2001; King et al. 2012; Brown et al. 2015; Zerbini et al. 2017). Typical of the majority of bipartite begomoviruses native to the New World, the DNA-A component of CuLCrV encodes six genes, two in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes has not been investigated for CuLCrV.

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682

Cucumis melo (Muskmelon or Cantaloupe)

Cucurbit yellow stunting disorder virus Taxonomic position Genus: Crinivirus

(CYSDV)

Family: Closteroviridae

Geographical distribution CYSDV was initially discovered in the United Arab Emirates in 1982 (Hassan and Duffus 1991). CYSDV infection in Cucumis melo plants was reported from United Arab Emirates, Spain, Portugal, Tunisia, Asia, Sudan, Lebanon, the desert southwest of the USA, Mexico, and Morocco (Livieratos et al. 1999; Desbiez et al. 2000; Kao et al. 2000; Louro et al. 2000; Sinclair and Crosby 2002; CABI/ EPPO 2004; Brown et al. 2007; Kuo et al. 2007; Wintermantel et al. 2007; Yakoubi et al. 2007; Liu et al. 2010; Tzanetakis et al. 2013; Abrahamian and Abou-Jawdah 2014). Symptoms and host(s) The virus-infected muskmelon plants express initial symptoms such as interveinal chlorosis and green spots on the oldest leaves. Severe symptoms include complete leaf yellowing (except veins) and overall brittleness. Plants are severely stunted and fruit quality is highly affected causing economic losses. CYSDV naturally infects certain hosts belonging to Amaranthaceae, Brassicaceae, Chenopodiaceae, Compositae, Euphorbiaceae, Fabaceae, Malvaceae, Portulacaceae, and Solanaceae. The wide natural host range of this virus was also confirmed through whitefly vector transmission (Wintermantel et al. 2009). Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a semi-persistent manner, with biotype B being a more effective vector than by biotype A (Celix et al. 1996; Wisler et al. 1998; McGrath 2004; Navas-Castillo et al. 2014). However, biotype Q transmits as efficiently as biotype B (Berdiales et al. 1999). Acquisition periods of 18 h or more and inoculation periods of 24 h or more are necessary for transmission rates of CYSDVof over 80%. The virus is not mechanically sap-transmissible. The virus is not seed-transmitted. Virion properties and genome The virions are non-enveloped, bipartite filamentous particles about 650–850 nm and 700–900 nm in length, and 10–13 nm in diameter (Liu et al. 2010). The genome is a positive-sense, single-stranded RNA (Celix et al. 1996). The genome is divided into two molecules RNA1 of 9123 nt (NC_004809) and RNA2 of 7976 nt (NC_004810) (Aguilar et al. 2003). Virus particles are encapsidated mainly by a major coat protein of 28 kDa (Livieratos et al. 1999; Livieratos and Coutts 2002; Coutts and Livieratos 2003; Kreuze 2011).

Eggplant mottled dwarf nucleorhabdovirus Taxonomic position Genus: Nucleorhabdovirus

(EMDV)

Family: Rhabdoviridae

EMDV infection in plants of Cucumis melo was reported from Italy (Ciuffo et al. 1999). The virusinfected muskmelon plants exhibit symptoms of stunting, short internodes, fruit deformation, and vein yellowing. The virus is transmitted by a leafhopper vector, Agallia vorobjevi, in a persistent,

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propagative manner, and also by mechanical sap-inoculation to solanaceous hosts. For more details of EMDV, refer to Solanum melongena.

Groundnut bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(GBNV)

C Family: Tospoviridae

GBNV infection in plants of Cucumis melo was reported from India (Holkar et al. 2013, 2017). The virus-infected muskmelon plants exhibit symptoms of mottling, yellowing of leaves, necrosis of shoot buds, and stunting of plants. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of GBNV, refer to Arachis hypogaea.

Kyuri green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(KGMMV)

Family: Virgaviridae

KGMMV infection in plants of Cucumis melo was reported from Indonesia (Francki et al. 1986; Daryono et al. 2005). The virus-infected muskmelon plants exhibit fruit deformation and mosaic symptoms. The virus is transmitted by means not involving a vector. The virus is transmissible by mechanical sap-inoculation, and also by contact between plants. For more details of KGMMV, refer to Cucumis sativus.

Lettuce infectious yellows virus Taxonomic position Genus: Crinivirus

(LIYV)

Family: Closteroviridae

LIYV was first reported in plants of Cucumis melo from Arizona and California (USA) in 1980–1981 and subsequently from Mexico (Brown and Nelson 1986; Brown and Poulos 1989; Halliwell and Johnson 1992). The virus-infected muskmelon plants exhibit chlorosis accompanied by a green mosaic. Symptoms develop primarily on older leaves. The plants are stunted and fruits usually do not ripen completely. The virus is transmitted by the whitefly vector Bemisia tabaci in a semi-persistent manner. Whiteflies acquire the virus in several minutes and transmit it for a period of 4–7 days. The virus is not seed-borne or not transmitted mechanically. For more details of LIYV, refer to Lactuca sativa.

Melon aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(MABYV)

Family: Luteoviridae

684

Cucumis melo (Muskmelon or Cantaloupe)

Geographical distribution MABYV infection in plants of Cucumis melo was reported from China (Xiang et al. 2008b). Symptoms and host(s) The virus-infected muskmelon plants exhibit yellowing symptoms on the leaves. Transmission The virus is transmitted by an aphid vector, Aphis gossypii, in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. Virion properties and genome The genome consists of a single molecule of infectious, linear, positive-sense ssRNA of 5674 nt (EU000534). The 30 terminus has neither a poly (A) tract nor a tRNA-like structure. The 50 terminus has a genome-linked protein (VPg) (of about 17 kDa). The genome contains six open reading frames (ORFs) (Xiang et al. 2008b).

Melon chlorotic leaf curl virus

(MCLCuV)

Synonyms Squash yellow mild mottle virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution MCLCuV infection in plants of Cucumis melo was reported from Zacapa Valley, Guatemala, and Costa Rica (Brown et al. 2001; Rosario et al. 2015). Symptoms and host(s) The virus-infected muskmelon plants exhibit patchy foliar chlorosis and leaf curl symptoms. The virus infection results in reduced fruit set. Transmission The transmission of the MCLCuV has not been investigated. It is likely that, in common with other begomoviruses, MCLCuV is transmitted by the whitefly vector Bemisia tabaci in a circulative, nonpropagative manner (Brown et al. 2001). Experimentally the virus has been introduced into common bean (Phaseolus vulgaris), cantaloupe (Cucumis melo), cucumber (Cucumis pepo), Datura stramonium, Nicotiana benthamiana, pepper (Capsicum annuum), pumpkin (Cucurbita maxima), sweet basil (Ocimum basilcum), and watermelon (Citrullus lanatus) by Agrobacterium-mediated inoculation of cloned virus genomes (Idris et al. 2008). Symptoms were leaf curling, foliar mottling, and stunting. Virion properties and genome The virion structure of MCLCuV has not been investigated. In common with all geminiviruses, the virions of MCLCuV are likely geminate (twinned quasi-icosahedra). MCLCuV is typical of bipartite begomoviruses native to the New World. The genome consists of two circular, single-stranded DNA components. DNA-A contains 2662 nt (NC_004732; AY064391 =

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NC_003865; AF325497, KC153490) and DNA-B of 2638 nt (NC_028138; AF440790 = NC_003860; AF325498) (Briddon 2001; Brown et al. 2001, 2015; Idris et al. 2008; Zerbini et al. 2017). Typical of the majority of bipartite begomoviruses native to the New World, the DNA-A component of MCLCuV encodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes has not been investigated for MCLCuV.

Melon chlorotic mosaic virus Taxonomic position Genus: Begomovirus

(MClMV)

Family: Geminiviridae

Geographical distribution MClMV infection in plants of Cucumis melo was reported from Venezuela (Ramirez et al. 2004; Romay et al. 2014, 2015). Symptoms and host(s) The virus-infected muskmelon plants exhibit yellow mosaic symptoms on the leaves. The natural host range of this virus is restricted to cucurbit species including watermelon (Citrullus lanatus), muskmelon (Cucumis melo), wild melon (Cucumis melo var. agrestis), cucumber (Cucumis sativus), Cucumis anguria, Cucumis dipsaceus, and Cucurbita moschata. Infected cucurbits show leaf curling and foliar chlorosis. Transmission The transmission of MeClMV has not been investigated. It is likely that, in common with other begomoviruses, MeClMV is transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. The virus is mechanically sap-transmissible to Cucumis melo, Nicotiana benthamiana, and Nicotiana clevelandii (Romay et al. 2015). Virion properties and genome The virion structure of MeClMV has not been investigated. In common with all geminiviruses, the virions of MeClMV are likely geminate (twinned quasi-icosahedra). MeClMV is typical of bipartite begomoviruses native to the New World. The genome consists of two circular, single-stranded DNA components. DNA-A contains 2628 nt (HM163576 = NC_014380) and DNA-B of 2607 nt (HM163577 = NC_014381) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). MeCMV has been associated with the satellite Melon chlorotic mosaic alphasatellite (MeCMA) (Romay et al. 2015). Typical of the majority of bipartite begomoviruses native to the New World, the DNA-A component of MeClMV encodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes has not been investigated for MeClMV.

Melon mild mottle virus Taxonomic position Genus: Nepovirus

(MMMoV)

Family: Secoviridae

C

686

Cucumis melo (Muskmelon or Cantaloupe)

Geographical distribution MMMoV infection in plants of Cucumis melo was reported from Japan (Tomitaka et al. 2011). Symptoms and host(s) The virus induced a mild mottle in muskmelon, but symptoms were only visible early in infection. The experimental host range of this virus is very wide, including at least 19 species in five families, with most showing even in systemically-infected leaves (Tomitaka et al. 2011). Transmission No vector is known for this virus. Seed and/or pollen transmission is possible but has not been confirmed experimentally. Absence of detectable nematodes from affected fields suggested seed transmission (Tomitaka et al. 2011). Virion properties and genome The virions are isometric, non-enveloped of two types but similar in size, 25–30 nm in diameter, and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is bipartite, linear positive-sense, and single-stranded RNA. RNA1 consists of 7727 nt (AB518485) and RNA2 is 3854 nt (AB518486) (Sanfacon et al. 2009; Dunez and Le Gall 2011; Tomitaka et al. 2011; Sanfacon 2015; Thompson et al. 2017).

Melon necrotic spot virus Taxonomic position Genus: Gammacarmovirus

(MNSV)

Family: Tombusviridae

Geographical distribution MNSV infection was first reported in plants of Cucumis melo from Japan by Kishi (1966). The virus spreads in Italy, Iran, Tunisia, China, Japan, the Netherlands, the UK, Guatemala, Mexico, Panama, Honduras, Uruguay, Spain, India, and the USA (Komuro 1971; Yoshida et al. 1980b; Avgelis 1990; Matsuo et al. 1991; Tomassoli and Barba 2000; Choi et al. 2003; Gosalvez et al. 2003; Kubo et al. 2005; Herrera et al. 2006; Safaeezadeh 2007; De Cara et al. 2008; Gu et al. 2008; Ohki et al. 2008; Yakoubi et al. 2008; CABI/EPPO 2010; Herrera-Vasquez et al. 2010; Nagateja and Gopinath unpublished JX879088). Symptoms and host(s) The virus-infected muskmelon plants exhibit necrosis of leaves, stems, branches, and fruits, leading to severe deterioration in fruit quality and serious economic damage. Some cultivars exhibit only necrotic spotting symptoms. The natural host ranges of this virus are melon (Cucumis melo), cucumber (Cucumis sativus), and watermelon (Citrullus lanatus) (Hibi and Furuki 1985). Transmission The virus is transmitted by the fungal vector Olpidium bornovanus (De Cara et al. 2008; HerreraVasquez et al. 2010). The virus is transmissible by mechanical sap-inoculation to less than three plant families and also transmissible by contact between plants. The virus is transmitted by seed to the extent of 10–40% in melon (Campbell et al. 1996; Herrera-Vasquez et al. 2009).

Cucumis melo (Muskmelon or Cantaloupe)

687

Virion properties and genome The virions are isometric, 30 nm in diameter, and have T = 3 icosahedral symmetry. The genome consists of a single molecule of linear positive-sense ssRNA of 4266 nt (M29671 = NC_001504) (Furuki et al. 1980; Riviere and Rochon 1990; Diaz et al. 2003). Several MNSV isolates from melon have been sequenced, including MNSV-Dutch, MNSV-NH and MNSV-NK, MNSV-YS and MNSVYK, and MNSV-chi (Gonzalez-Garza et al. 1979; Ohshima et al. 2000).

Melon rugose mosaic virus Taxonomic position Genus: Tymovirus

(MRMV)

Family: Tymoviridae

MRMV infection in plants of Cucumis melo was reported from Yemen and Sudan (Jones et al. 1986; Mahgoub et al. 1997). The virus-infected muskmelon plants exhibit severe mosaic symptoms. The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sapinoculation. The virus is seed-transmitted in Cucumis melo up to 0.9% (Mahgoub et al. 1997). For more details of MRMV, refer to Citrullus lanatus.

Melon severe mosaic orthotospovirus Taxonomic position Genus: Orthotospovirus

(MSMV)

Family: Tospoviridae

Geographical distribution MSMV infection in plants of Cucumis melo was reported from Mexico (Ciuffo et al. 2009, 2017).

Symptoms and host(s) The virus-infected muskmelon plants show leaf mosaic, blistering, or deformation, as well as fruit splitting or necrotic lesions on mature fruit.

Transmission The virus is transmitted by a thrips vector Frankliniella occidentalis in a persistent-propagative manner. The virus is mechanically sap-transmissible to a number of herbaceous hosts including N. benthamiana, N. megalosiphon, Datura stramonium, and N clevelandii and caused deformation, mosaic, or necrosis. The virus infection caused also chlorotic spots on Emilia sonchifolia plants.

Virion properties and genome The virions are spherical or pleiomorphic, and membrane bound 80-120 nm in diameter. The genome comprises three unique molecules of negative or ambisense ssRNA species: small (S) RNA of 3283nt (KX698422 = NC_033832); medium (M) RNA of 4873nt (KX698423 = NC_033833); and large (L) RNA of 8911nt(KX698424 = NC_033834) (Ciuffo et al. 2017).

C

688

Cucumis melo (Muskmelon or Cantaloupe)

Melon variegation virus

(MVV)

Taxonomic position MVV is a tentative member of the family Rhabdoviridae. Geographical distribution MVV infection was first reported in plants of Cucumis melo from Spain (Rubio-Huertos and PenaIglesias 1973). The virus spreads in Spain. Symptoms and host(s) The virus-infected muskmelon plants exhibit symptoms of stunting WITH chlorosis of leaves and fruits. Transmission There is no known vector for this virus. Virion properties and genome The virions are rhabdo- or bullet-shaped with a clear modal length of 320 nm and 60 nm wide.

Melon vein-banding mosaic virus

(MVbMV)

Taxonomic position MVbMV is a tentative member of the genus Potyvirus and family Potyviridae Geographical distribution MVbMV infection was first reported in plants of Cucumis melo from Taiwan by Huang et al. (1993). The virus spreads in Taiwan. Symptoms and host(s) The virus-infected muskmelon plants exhibit mosaic or vein-banding symptoms. Transmission The virus is transmitted by an aphid vector, Aphis gossypii, in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are flexuous filaments rods of c.755 nm long. The genome consists of a single molecule of positive-sense single-stranded RNA of c.9.5–10 kb. A partial genome sequence of 1108 nt is available (HQ423412) (Huang et al. 1993; Wylie et al. 2017).

Melon yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(MYSV)

Family: Tospoviridae

Cucumis melo (Muskmelon or Cantaloupe)

689

Geographical distribution MYSV infection in plants of Cucumis melo was reported from Ecuador, Japan, Thailand, and China (Kato et al. 1999, 2000; Peng et al. 2011; Chen et al. 2012; Gu et al. 2012; Quito-Avila et al. 2014). Symptoms and host(s) The virus-infected muskmelon plants exhibit symptoms of vein-clearing and chlorotic spots on newly developed leaves. These leaves showed yellowing and necrotic spots as they aged. If infection occurred early, mosaic symptoms form on fruit. Infection had a deleterious effect on the visual appeal and the taste of fruit (Kato et al. 1999; Okuda et al. 2002; Gu et al. 2012). Transmission The virus is transmitted by a thrips vector, Thrips palmi, in a persistent, propagative manner (Kato et al. 2000). The virus is mechanically sap-transmissible. Virion properties and genome The virions are spherical or pleomorphic and 80–120 nm in diameter (King et al. 2012). The virus has a genome consisting of three single-stranded RNA molecules, namely the large (L) 8918 nt [AB061774 = NC_008306], medium (M) 4815 nt [AB061773 = NC_008307] and small (S) 3232 nt [AB038343 = NC_008300] RNAs (Kato and Hanada 2000; Kato et al., 2000; Okuda et al., 2004, 2006; Chiemsombat et al., 2008; Chen et al., 2012).

Melon yellowing-associated virus Taxonomic position Genus: Carlavirus

(MYaV)

Family: Betaflexiviridae

Geographical distribution MYaV infection in plants of Cucumis melo was reported from Brazil (Nagata et al. 2003, 2005; Lima et al. 2009; Costa et al. 2017). Symptoms and host(s) The virus-infected muskmelon plants exhibit symptoms of yellowing and mottling on older leaves. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, B biotype in a semi-persistent manner (Nagata et al. 2005). The virus is also mechanically sap-transmissible. Virion properties and genome The virions are filamentous particles 600–700 nm in length. The genome consists of a single molecule of linear positive-sense ssRNA. The virus was initially identified as a distinct carlavirus based on the 30 terminal genome sequence of 1612 nt (AY373028) and contains complete coat protein and nucleic acidbinding protein (Nagata et al. 2005). Later, the 30 terminal genome sequence was extended up to 3137 nt (AB510477) that included the triple gene block region (Adams et al. 2004; Nagata et al. 2010; Costa et al. 2017).

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690

Ourmia melon virus Taxonomic position Genus: Ourmiavirus

Cucumis melo (Muskmelon or Cantaloupe)

(OuMV)

Family: Botourmiaviridae

Geographical distribution OuMV infection was first reported in plants of Cucumis melo from Ourmia, northern Iran, by Lisa et al. (1988). The virus spreads in Iran (Gholamalizadeh et al. 2008). Symptoms and host(s) The virus-infected muskmelon plants exhibit symptoms of chlorotic spots and irregular ringspots and puckering. Transmission No vector has been identified for this virus. The virus is mechanically sap-transmissible to a number of dicotyledon plants. Virion properties and genome The virions are bullet-shaped with conical ends, 18.5 nm in mean width, parallel-sided, and of several discrete lengths, 30 nm and 37 nm being the shortest and commonest lengths. The genome is a linear, single-stranded, positive-sense RNA of three sizes. RNA1 consists of 2814 nt (EU770623 = NC_011068), RNA2 of 1064 nt (EU770624 = NC_011069), and RNA3 is 974 nt (EU770625 = NC_011070) (Marzachi et al. 1992; Accotto et al. 1997; Rastgou et al. 2009; Milne and Accotto 2011; Turina et al. 2017).

Papaya ringspot virus Taxonomic position Genus: Potyvirus

(PRSV)

Family: Potyviridae

PRSV infection in plants of Cucumis melo was reported from the USA and Spain (Rivera et al. 1991; Grafton-Cardwell et al. 1996; Luis-Arteaga et al. 1998; Romay et al. 2014; Tantiwanich et al. 2014; Nagendran et al. 2017). The virus-infected muskmelon plants exhibit mosaic and mottling symptoms resulting in vine decline and reduction in fruit yield and quality. The virus is transmitted by the aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PRSV, refer to Carica papaya.

Squash chlorotic leaf spot virus Taxonomic position Genus: Torradovirus

(SCLSV)

Family: Secoviridae

SCLSV infection in plants of Cucumis melo was reported from Sudan (Lecoq et al. 2016). The virusinfected plants exhibit chlorotic leaf spot symptoms. The virus is transmitted by the greenhouse whitefly (Trialeurodes vaporariorum) and silverleaf whitefly (Bemisia tabaci) in a semi-persistent manner. The virus is also mechanically transmissible. For more details of SCLSV refer, to Cucurbita pepo.

Cucumis melo (Muskmelon or Cantaloupe)

Squash leaf curl virus Taxonomic position Genus: Begomovirus

691

(SLCuV)

Family: Geminiviridae

SLCuV infection in plants of Cucumis melo was reported from Israel, Lebanon, and California (Lazarowitz and Lazdins 1991; Kuo et al. 2007; Ali-Shtayeh et al. 2010; Sufrin-Ringwald and Lapidot 2011; Sobh et al. 2012). The virus-infected muskmelon plants initially showed symptoms of mild leaf curling, some chlorotic lesions, and enations on the affected leaves. However within 2–3 weeks of the first appearance of symptoms, the plants recovered and symptoms completely disappeared. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of SLCuV, refer to Cucurbita pepo.

Squash mosaic virus Taxonomic position Genus: Comovirus

(SqMV)

Family: Secoviridae

SqMV infection in plants of Cucumis melo was reported from Brazil, Greece, Japan, Turkey, Iran, and Morrocco (Kemp et al. 1972; Yoshida et al. 1980a; Lockhart et al. 1982; Izadpanah 1987; Avgelis and Katis 1989; Caglar et al. 2004; Lima et al. 2012; Romay et al. 2014). The virus-infected muskmelon plants exhibit symptoms of leaves including green vein-banding, mosaic, mottling, ring patterns of leaves, leaf deformations and enations and mottling symptoms on fruits. The virus is transmitted by beetle vectors in a non-persistent manner, and also by mechanical sap-inoculation. High rate (up to 100%) of seed transmission of the virus was reported in Cucumis melo (Kemp et al. 1972; Alvarez and Campbell 1978; Franken et al. 1990; Lestari and Nurhayati 2014). For more details of SqMV, refer to Cucurbita pepo.

Squash vein yellowing virus Taxonomic position Genus: Ipomovirus

(SqVYV)

Family: Potyviridae

SqVYV infection in plants of Cucumis melo var. dudaim was reported from Florida (Adkins et al. 2009). The virus-infected muskmelon plants exhibit chlorosis of lower leaves. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a non-persistent manner, and also by mechanical sapinoculation to other cucurbitaceous species. For more details of SqVYV, refer to Cucurbita pepo.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Cucumis melo was reported from North America, Australia, Nigeria, Iran, Japan, and the Netherlands (Yoshida et al. 1980b). The virus-infected muskmelon plants exhibit very

C

692

Cucumis melo (Muskmelon or Cantaloupe)

bright yellow mosaic, ringspots, necrosis, and distortion symptoms. The virus is transmitted by the dagger nematode vector, Xiphinema americanum, in a non-persistent manner, and also by mechanical sap-inoculation. The virus is transmitted through seed of Cucumis melo to the extent of 3–7% (McLean 1962). The virus is not transmitted by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Cucumis melo was reported from Iran, Tunisia, Thailand, and Taiwan (Ito et al. 2008; Chang et al. 2010; Mnari-Hattab et al. 2015; Yazdani-Khameneh et al. 2016; Samretwanich et al. 2000a, b). The virus-infected muskmelon plants exhibit severe yellow leaf symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative nonpropagative manner. The virus is mechanically transmissible (Chang et al. 2010). The virus is grafttransmissible. For more details of ToLCNDV, refer to Solanum lycopersicum.

Tomato leaf curl Palampur virus Taxonomic position Genus: Begomovirus

(ToLCPalV)

Family: Geminiviridae

ToLCPalV infection in plants of Cucumis melo was reported from Iran (Heydarnejad et al. 2013). The virus-infected muskmelon plants exhibit leaf curl symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is graft-transmissible. For more details of ToLCPalV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Cucumis melo was reported from Japan (Yoshida et al. 1980b). The virusinfected muskmelon plants exhibit yellow mosaic symptoms and prominent ringspots on the discolored fruits. Leaves are reduced in size and plants have shortened internodes. The virus is transmitted by a nematode vector, Xiphinema americanum, in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and by but seed transmission was not detected in C. melo (Yoshida et al. 1980b). For more details of ToRSV, refer to Solanum lycopersicum.

Watermelon bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(WBNV)

Cucumis melo (Muskmelon or Cantaloupe)

693

WBNV infection in plants of Cucumis melo was reported from India (Holkar et al. 2017). The virusinfected muskmelon plants exhibit bud necrosis, mottling, and veinal necrosis symptoms. The virus is transmitted by the thrips vectors in a persistent-propagative manner, and also by mechanical sapinoculation. The virus is not transmitted by contact between plants. For more details of WBNV, refer to Citrullus lanatus.

C Watermelon chlorotic stunt virus Taxonomic position Genus: Begomovirus

(WmCSV)

Family: Geminiviridae

WmCSV infection in plants of Cucumis melo was reported from Sudan, Israel, and Lebanon (Dafalla et al. 1998; Antignus et al. 2003; Sufrin-Ringwald and Lapidot 2011; Samsatly et al. 2012). The virusinfected muskmelon plants usually developed initial symptoms as yellow pinpoints that turned into large chlorotic lesions with time. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by mechanical inoculation but transmissible by grafting. For more details of WmCSV, refer to Citrullus lanatus.

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV infection in plants of Cucumis melo was reported from Brazil, California, Japan, Argentina, Italy and China (Yoshida et al. 1980b; Al-Musa and Mansour 1982; Grafton-Cardwell et al. 1996; LuisArteaga et al. 1998; Meneghini et al. 2007; Lima et al. 2012; Trkulja et al. 2014; Perotto et al. 2016; Wang et al. 2017). The virus-infected muskmelon plants exhibit severe vein-banding, green mosaic, deformation, and plant stunting. The virus is aphid transmitted in a non-persistent manner (Garzo et al. 2004). The virus is mechanically sap-transmissible, and its host range was restricted mostly to Cucurbitaceae, Leguminosae, and Chenopodiaceae. For more details of WMV, refer to Citrullus lanatus.

Watermelon silver mottle orthotospovirus Taxonomic position Genus: Orthotospovirus

(WSMoV)

Family: Tospoviridae

WSMoV infection in plants of Cucumis melo was reported from Japan and Taiwan (Okuda et al. 2002). The virus-infected muskmelon plants exhibit symptoms of foliar mottling, plant stunting, upright growth of branches, and tip blight. Silver mottling occurred on leaves and chlorotic mottling on fruits. The virus is transmitted by a thrips vector, Thrips palmi, in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of WSMoV, refer to Citrullus lanatus.

694

Cucumis melo (Muskmelon or Cantaloupe)

Zucchini yellow fleck virus Taxonomic position Genus: Potyvirus

(ZYFV)

Family: Potyviridae

ZYFV infection in plants of Cucumis melo was reported from Israel, Italy, and some other Mediterranean countries (Antignus et al. 1995; Desbiez et al. 2007; Tomassoli et al. 2010). The virus-infected muskmelon plants exhibit systemic mosaic or necrotic spots, and even complete necrosis of plants. The virus is transmitted by the aphid vector, Myzus persicae, in a non-persistent manner, and also is transmissible through mechanical sap-inoculation. For more details of ZYFV, refer to Cucurbita pepo.

Zucchini yellow mosaic virus Taxonomic position Genus: Potyvirus

(ZYMV)

Family: Potyviridae

ZYMV infection in plants of Cucumis melo was reported from southeastern California desert valleys, Italy, Brazil, Spain, China, and India (Lecoq and Pitrat 1984; Nameth et al. 1985; Orozco-Santos et al. 1994; Grafton-Cardwell et al. 1996; Luis-Arteaga et al. 1998; Singh et al. 2004; Malik et al. 2006, 2010; Lima et al. 2012; Sharma and Gaur 2015; Wang and Li 2017; Wang et al. 2017). Early symptoms on leaves of ZYMV-infected muskmelon are vein-clearing and yellow mosaic. Leaves are subsequently reduced in size, deformed, often with serrated edges and dark green blisters or enations, and contrasting with the yellow or light green color of the rest of the lamina. Branches develop short internodes and usually exhibit an erect habit. Discoloration, necrotic cracks, and raised patches are observed on fruits. The virus is transmitted by aphid vectors such as Myzus persicae and Aphis gossypii in a nonpersistent manner, and also by mechanical sap-inoculation (Garzo et al. 2004). Seeds are deformed and have low germination rates (Provvidenti and Robinson 1987). For more details of ZYMV, refer to Cucurbita pepo.

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In: Encyclopedia of Life Sciences. John Wiley and Sons Ltd, Chichester. http://www.els.net. https://doi.org/10.1002/ 9780470015902.a0000764.pub3 Sanfacon H, Wellink J, Le Gall O, Karasev A, van der Vlugt R, Wetzel T (2009) Secoviridae: a proposed family of plant viruses within the order Picornavirales that combines the families Sequiviridae and Comoviridae, the unassigned genera Cheravirus and Sadwavirus, and the proposed genus Torradovirus. Arch Virol 154(5):899–907 Sharma P, Gaur RK (2015) First report of Zucchini yellow mosaic virus on muskmelon in India. J Plant Pathol 97:541–551 Sharma OP, Khatri HL, Bansal RD,Komal HS (1984) A new strain of Cucumber mosaic virus causing mosaic disease of muskmelon. J Phytopath 109:332–340 Sinclair JW, Crosby KM (2002) A review of Cucurbit yellow stunting disorder virus (CYSDV) – a “new” virus affecting melons in the lower Rio Grande Valley. Subtrop Plant Sci 54:54–58 Singh SJ, Verma R, Singn RK, Prakash S (2004) First report of natural occurrece of Zucchini yellow mosaic potyvirus (ZYMV) on musk melon. Indian Plant Pathol 22:134–135 Sobh H, Samsatly J, Jawhari M, Najjar C, Haidar A, Abou-Jawdah Y (2012) First report of Squash leaf curl virus in cucurbits in Lebanon. Plant Dis 96(8):1231 Sufrin-Ringwald T, Lapidot M (2011) Characterization of a synergistic interaction between two cucurbit-infecting begomoviruses: squash leaf curl virus and Watermelon chlorotic stunt virus. Phytopathology 101:281–289 Tantiwanich Y, Baker CA, Turechek WW, Adkins S (2014) Detection of Papaya ringspot virus type W infecting the cucurbit weed Cucumis melo var. dudaim in Florida. Plant Health Progress. https://doi.org/10.1094/PHP-BR-13-0126 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J GenVirol 98:529–531 Tian T, Posis K, Maroon-Lango CJ, Mavrodieva V, Haymes S, Pitmal TL, Falk BW (2014) First report of Cucumber green mottle mosaic virus on Melon in the United States. Plant Dis 98:1163–1164 Tobias I, Maat DZ, Huttinga H (1982) Two Hungarian isolates of cucumber mosaic virus from sweet pepper (Capsicum annuum) and melon (Cucumis melo): identification and antiserum preparation. Neth J Plant Pathol 88:171–183

Cucumis melo (Muskmelon or Cantaloupe)

701

Tomassoli L, Barba M (2000) Occurrence of Melon necrotic spot carmovirus in Italy. EPPO Bull 30:279–280 Tomassoli L, Meneghini M (2007) First report of Cucurbit aphid-borne yellows virus in Italy. Plant Pathol 56:720 Tomassoli L, Lumia V, Siddu GF, Barba M (2003) Yellowing disease of melon in Sardinia (Italy) caused by Beet pseudoyellows virus. J Plant Pathol 85:59–61 Tomassoli L, Tiberini A, Meneghini M (2010) Zucchini yellow fleck virus is an emergent virus on melon in Sisily (Italy). J Phytopathol 158:314–316 Tomitaka Y, Usugi T, Yasuda F, Okayama H, Tsuda S (2011) A novel member of the genus Nepovirus isolated from Cucumis melo in Japan. Phytopathology 101:316–322 Trkulja V, Kovacic D, Curkovic B, Vucurovic A, Stankovic I, Bulajic A, Krstic B (2013) First report of Cucumber mosaic virus on Melon in Bosnia and Herzegovina. Plant Dis 97:1124–1125 Trkulja V, Vasic J, Vukovic B, Stankovic I, Vucurovic A, Bulajic A, Krstic B (2014) First report of Watermelon mosaic virus infecting Melon and Watermelon in Bosnia and Herzegovina. Plant Dis 98:1749 Turina M, Hillman BI, Izadpanah K, Rastgou M, Rosa C, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Ourmiavirus. J GenVirol 98:129–130 Tzanetakis IE, Martin RR, Wintermantel WM (2013) Epidemiology of criniviruses: an emerging problem in world agriculture. Front Microbiol 4:119 Vani S, Varma A (1993) Properties of Cucumber green mottle mosaic virus isolated from water of river Jamuna. Indian Phytopath 46:118–122 Wang D, Li G (2017) Biological and molecular characterization of Zucchini yellow mosaic virus isolates infecting melon in Xinjiang, China. Can J Plant Pathol 39:48–59 Wang D, Li G, Du SS (2017) Occurrence of viruses infecting melon in Xinjiang of China and molecular characterization of Watermelon mosaic virus isolates. Eur J Plant Pathol 147:919–931 Wintermantel WM, Brown JK, Gilbertson RL (2007) Wide spread emergence of Cucurbit yellow stunting disorder virus (CYSDV) in the southwestern desert melon production region of the United States and Mexico. Phytopathology 97:S124 Wintermantel WM, Hladky LL, Cortez AA, Natwick ET (2009) A new expanded host range of Cucurbit yellow stunting disorder virus includes three agricultural crops. Plant Dis 93:685–690 Wisler GC, Dufus JE, Liu H-Y, Li RH (1998) Ecology and epidemiology of whitefly-transmitted closteroviruses. Plant Dis 82:270–280 Wu HJ, Qin BX, Chen HY, Peng B, Cai JH, Gu QS (2011) The rate of seed contamination and transmission of Cucumber green mottle mosaic virus in watermelon and melon. Sci Agric Sin 44:1527–1532 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J GenVirol 98:352–354 Xiang HY, Shang QX, Han CG, Li DW, Yu JL (2008a) First report on the occurrence of Cucurbit aphid-borne yellows virus on nine cucurbitaceous species in China. Plant Pathol 57:390 Xiang HY, Shang QX, Han CG, Li DW, Yu JL (2008b) Complete sequence analysis reveals two distinct poleroviruses infecting cucurbits in China. Arch Virol 153:1155–1160 Yakoubi S, Desbiez C, Fakhfakh H, Wipf-Scheibel C, Marrakchi M, Lecoq H (2007) Occurrence of Cucurbit yellow stunting disorder virus and Cucumber vein yellowing virus in Tunisia. J Plant Pathol 89(3):417–420 Yakoubi S, Desbiez C, Fakhfakh H, Wipf-Scheibel C, Marrakchi M, Lecoq H (2008) First report of Melon necrotic spot virus on melon in Tunisia. Plant Pathol 57:386 Yamashita S, Doi Y, Yora K, Yoshino M (1979) Cucumber yellows virus: Its transmission by the greenhouse whitefly, Trialeurodes vaporariorum (Westwood), and the yellowing disease of cucumber and muskmelon caused by the virus. Ann Phytopathol Soc Jpn 45:484–496 Yardimci N, Ozgonen H (2007) First report of Cucurbit aphid-borne yellows virus in Turkey. Aust Plant Dis Notes 2:59 Yazdani-Khameneh S, Aboutorabi S, Shoori M, Aghazadeh A, Jahanshahi P, Golnaraghi A, Maleki M (2016) Natural occurrence of tomato leaf curl New Delhi virus in Iranian cucurbit crops. Plant Pathol J 32:201–208 Yilmaz MA, Ozaslan M, Ozaslan D (1989) Cucumber vein yellowing virus in Cucurbitaceae in Turkey. Plant Dis 73:610 Yoshida K, Goto T, Nemoto M (1980a) Squash mosaic virus isolated from melon (Cucumis melo L.) in Hokkaido. Ann Phytopathol Soc Jpn 46:349–356 Yoshida K, Goto T, Nemoto M, Tsuchizaki T (1980b) Five viruses isolated from melon (Cucumis melo L.) in Hokkaido. Ann Phytopathol Soc Jpn 46:339–348 Zeng R, Dai FM, Chen WJ, Lu JP (2011) First report of Cucurbit chlorotic yellows virus infecting melon in China. Plant Dis 95:354 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J GenVirol 98:131–133 Zhang YJ, Li GF, Li MF (2009) Occurrence of Cucumber green mottle mosaic virus on cucurbitaceous plants in China. Plant Dis 93:200 Zindovic J, Manglli A, Hrncic S, Radonjic S, Perovic T, Tomassoli L (2017) First report of Cucurbit aphid-borne yellows virus affecting summer squash and melon in Montenegro. J Plant Pathol 99:299

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702

Cucumis melo var. agrestis (Wild melon)

Cucumis melo var. agrestis (Wild melon) Family: Cucurbitaceae

Medicinal

Wild melon banding virus Taxonomic position Genus: Potyvirus

(WMBV)

Family: Potyviridae

Geographical distribution WMBV infection in plants of Cucumis melo var. agrestis was reported from Sudan (Desbiez et al. 2017). Symptoms and host(s) The virus-infected wild melon plants exhibit mild vein banding and mosaic symptoms. Transmission The virus is transmitted by an aphid vector, Myzus persicae in a non-persistent manner. The virus is transmitted by mechanical sap-inoculation. The virus is not transmitted by contact between plants; not transmitted by seed; not transmitted by pollen. Virion properties and genome The virions are non-enveloped, flexuous filaments 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 10152 nt (KY623506 = NC_035458) (Revers and Garcia 2015; Desbiez et al. 2017; Wylie et al. 2017).

References Desbiez C, Wipf-Scheibel C, Millot P, Verdin E, Dafalla G, Lecoq H (2017) New species in the Papaya ringspot virus cluster: insights into the evolution of the PRSV lineage. Virus Res 241:88–94 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: potyviridae. J Gen Virol 98:352–354

Cucumis melo var. flexuosus (Snake melon) Family: Cucurbitaceae

Vegetable

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

Family: Luteoviridae

(CABYV)

Cucumis melo var. flexuosus (Snake melon)

703

CABYV infection in plants of Cucumis melo var. flexuosus was reported from Turkey (Yardimci and Ozgonen 2007). The virus-infected snake melon plants exhibit symptoms of initial chlorotic lesions followed by yellowing of whole leaves and thickening of older leaves. The virus is transmitted by aphid vectors, Myzus persicae, M. euphorbiae, and Aphis gossypii, in a circulative, non-propagative manner (Lecoq et al. 1992). The virus is not transmissible by mechanical sap-inoculation. For more details of CABYV, refer to Cucumis melo.

Cucurbit yellow stunting disorder virus Taxonomic position Genus: Crinivirus

(CYSDV)

Family: Closteroviridae

CYSDV infection in plants of Cucumis melo var. flexuosus was reported from UAE (Hassan and Duffus 1991; McGrath 2004). The virus-infected snake melon plants exhibit severe yellowing and stunting symptoms. The virus is transmitted by the whitefly vector, both the B and Q biotypes of B. tabaci, in a semi-persistent manner. The virus is not mechanically sap-transmissible. For more details of CYSDV, refer to Cucumis melo.

Groundnut bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(GBNV)

Family: Tospoviridae

GBNV infection in plants of Cucumis melo var. flexuosus was reported from India (Holkar et al. 2013). The virus-infected snake melon plants exhibit chlorotic ringspot symptoms on the leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of GBNV, refer to Arachis hypogaea.

Melon rugose mosaic virus Taxonomic position Genus: Tymovirus

(MRMV)

Family: Tymoviridae

MRMV infection in plants of Cucumis melo var. flexuosus was reported from Sudan (Mahgoub et al. 1997). The virus-infected snake melon plants exhibit severe mosaic, leaf deformations, and blisters on intermediate leaves, while young leaves exhibit milder symptoms. The virus is transmitted by beetle vectors in a semi-persistent manner, and also by mechanical sap-inoculation to less than three families. The virus is seed-transmitted in snake melon up to 3.8% (Mahgoub et al. 1997). For more details of MRMV, refer to Citrullus lanatus.

Snake melon asteroid mosaic virus

(SMAMV)

Taxonomic position SMAMV is a tentative member of the genus Sobemovirus and family Solemoviridaeand.

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704

Cucumis melo var. flexuosus (Snake melon)

Geographical distribution SMAMV infection in plants of Cucumis melo var. flexuosus was reported from Sudan (Lecoq et al. 2005, 2011). Symptoms and host(s) The virus-infected snake melon plants show severe symptoms of star-shaped chlorotic spots, mosaic, stunting, and leaf deformation on the young leaves and yellowing symptoms on the older leaves. Transmission The virus is transmitted by the red melon beetle (Aulacophora foveicollis). Minimum acquisition and inoculation access periods of 3 h and 2 h, respectively are required for transmission of the virus. The virus is mechanically transmissible and has a host range limited to a few cucurbit species. The virus is transmitted through seed at a low rate. Virion properties and genome The virions are isometric approximately 30 nm in diameter (Lecoq et al. 2011; Somera et al. 2015). The genome is presumed to be single-stranded RNA; only a short sequence fragment is available (HM450304).

Sudan watermelon mosaic virus Taxonomic position Genus: Potyvirus

(SuWMV)

Family: Potyviridae

Geographical distribution SuWMV infection in plants of Cucumis melo var. flexuosus was reported from Sudan (Desbiez et al. 2017). Symptoms and host(s) The virus-infected snake melon plants exhibit symptoms of mosaic and leaf deformations. Transmission The virus is transmitted by an aphid vector Myzus persicae in a non-persistent manner. The virus is transmitted by mechanical sap inoculation. The virus is not transmitted by contact between plants; not transmitted by seed; not transmitted by pollen. Virion properties and genome The virions are nonenveloped, flexuous filaments 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 10232 nt (KY623505 = NC_035459) (Revers and Garcia 2015; Desbiez et al. 2017; Wylie et al. 2017).

Tomato leaf curl Palampur virus Taxonomic position Genus: Begomovirus

(ToLCPalV)

Family: Geminiviridae

Cucumis metuliferus (Horned melon)

705

ToLCPalV infection in plants of Cucumis melo var. flexuosus was reported from India (Raj et al. 2015). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner, and also by grafting. For more details of ToLCPalV, refer to Solanum lycopersicum.

References

C

Desbiez C, Wipf-Scheibel C, Millot P, Verdin E, Dafalla G, Lecoq H (2017) New species in the Papaya ringspot virus cluster: insights into the evolution of the PRSV lineage. Virus Res 241:88–94 Hassan AA, Duffus JE (1991) A review of a yellowing and stunting disorder of cucurbits in the United Arab Emirates. Emir J Agric Sci 2:1–16 Holkar SK, Mandal B, Jain RK (2013) New ornamental and cucurbitaceous hosts of groundnut bud necrosis virus in India. Indian J Virol 24:146 Lecoq H, Bourdin D, Wipf-Scheibel C, Bon M, Lot H, Lemaire O, Herrbach E (1992) A new yellowing disease of cucurbits caused by a Luteovirus, Cucurbit aphid-borne yellows virus. Plant Pathol 41:749–761 Lecoq H, Dafalla G, Desbiez C, Delecolle B, Wipf-Scheibel C, Girard M (2005) Characterization of Snake Melon Asteroid Mosaic Virus, a tentative new sobemovirus infecting cucurbits. 2nd Joint Conference of the International Working Groups on Legume (IWGLV) and Vegetable Viruses (IWGVV), April 10–14 (2005), held at Florida, USA (Abstract) Lecoq H, Dafalla G, Delécolle B, Wipf-Scheibel C, Desbiez C (2011) Snake melon asteroid mosaic virus, a tentative new member of the genus Sobemovirus infecting cucurbits. Plant Dis 95:153–157 Mahgoub HA, Wipf-Scheibel C, Delecolle B, Pitrat M, Dafalla G, Lecoq H (1997) Melon rugose mosaic virus: characterization of an isolate from Sudan and seed transmission in melon. Plant Dis 81:656–660 McGrath MT (2004) Diseases of cucurbits and their management. In: Naqvi SAMH (ed) Diseases of fruits and vegetables: diagnosis and management, vol 1. Kluwer Publishers, Dordrecht, pp 455–510 Raj SK, Kumar S, Srivastava A (2015) Association of Tomato leaf curl Palampur virus with yellow mosaic disease of Armenian cucumber (Cucumis melo var. flexuoses) and wild melon (C. callosus var. agrestis) in India. Arch Phytopathol Plant Protect 48:751–759 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Somera M, Sarmiento C, Truve E (2015) Overview on Sobemoviruses and a proposal for the creation of the family Sobemoviridae. Viruses 7:3076–3115 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Yardimci N, Ozgonen H (2007) First report of Cucurbit aphid-borne yellows virus in Turkey. Aust Plant Dis Notes 2:59

Cucumis metuliferus (Horned melon) Family: Cucurbitaceae

Edible fruit

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV infection in plants of Cucumis metuliferus was reported from Hungary (Szathmary et al. 2008). The virus-infected horned melon plants exhibit symptoms of severe mosaic, deformations, and enations. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of WMV, refer to Citrullus lanatus.

706

Cucumis sativus (Cucumber)

References Szathmary E, Salamon P, Paksi AM, Palkovics L (2008) Characterization of some Hungarian Watermelon mosaic virus (WMV) isolates collected from watermelon (Citrullus lanatus [Thunbg.] Mats. et Nak.), melon (Cucumis melo L.) and kiwano (Cucumis metuliferus Naud.). The 3rd Conference of the International Working Group on Legume and Vegetable Viruses (IWGLVV), August 20th–23rd, 2008, p 69

Cucumis sativus (Cucumber) Family: Cucurbitaceae

Vegetable

Artichoke yellow ringspot virus Taxonomic position Genus: Nepovirus

(AYRSV)

Family: Secoviridae

AYRSV infection in plants of Cucumis sativus was reported from Crete (Avgelis and Vovlas 1989). The virus-infected cucumber plants exhibit symptoms of stunting and bright yellow mosaic of the leaves. No vector is known for this virus. The virus is mechanically sap-transmissible. For more details of AYRSV, refer to Cynara cardunculus var. scolymus.

Beet pseudoyellows virus

(BPYV)

Synonyms Cucumber chlorotic spot virus; Cucumber yellows virus Taxonomic position Genus: Crinivirus

Family: Closteroviridae

BPYV infection in plants of Cucumis sativus was reported from Greece, France, Japan, Spain, Bulgaria, and New Zealand (Yamashita et al. 1979; Zenbayashi et al. 1984; Woudt et al. 1993; Clover et al. 2002; Hartono et al. 2003; Boubourakas et al. 2006). The virus-infected cucumber plants exhibit symptoms which start as chlorotic spots on older leaves sometimes near the petiole. The leaves become completely yellow except for the veins which remain green. Older leaves are thickened and brittle when crushed. Generally young leaves remain normal, but in some cultivars a complete plant chlorosis is observed. The virus is transmitted by the greenhouse whitefly Trialeurodes vaporariorum in a semi-persistent manner (Wisler et al. 1998; Caciagli 2001). The virus is not transmissible by mechanical sap-inoculation, and also not by contact between plants. For more details of BPYV, refer to Beta vulgaris.

Chickpea chlorotic dwarf virus Taxonomic position Genus: Mastrevirus

(CpCDV)

Family: Geminiviridae

Cucumis sativus (Cucumber)

707

CpCDV infection in plants of Cucumis sativus was reported from Pakistan (Hameed et al. 2017). The virus-infected cucumber plants exhibit symptoms of leaf curling, mosaic, and necrotic spots. The virus is transmitted by leafhopper vectors in a persistent, circulative, and non-propagative manner. The virus is not transmissible by mechanical inoculation. For more details of CpCDV, refer to Cicer arietinum.

Cucumber Bulgarian latent virus

(CBLV)

Synonyms Cucumber Bulgarian virus Taxonomic position Genus: Tombusvirus

Family: Tombusviridae

Geographical distribution CBLV infection in plants of Cucumis sativus was first reported from Bulgaria (Kostova et al. 2003). The virus spreads in Bulgaria and Iran (Kostova et al. 2003; Menzel et al. 2015). Symptoms and host(s) The virus-infected cucumber plants do not exhibit any external symptoms. Transmission There is no known vector for this virus. The virus is mechanically sap-transmissible. Virion properties and genome The virions are isometric and icosahedral (T = 3), non-enveloped, and about 32–35 nm in diameter. The genome is a monopartite, linear, single-stranded, positive-sense RNA of 4576 nt (AY163842 = NC_004725) (White 2011; Menzel et al. 2015).

Cucumber cryptic virus

(CuCV)

Taxonomic position CuCV is a tentative member of the genus Alphacryptovirus. Geographical distribution CuCV infection in plants of Cucumis sativus was first reported from Germany (Jelkmann et al. 1988). Symptoms and host(s) The virus-infected cucumber plants do not exhibit any external symptoms. Transmission The virus is not mechanically sap-transmissible but transmitted through seed. Virion properties and genome The virions are isometric and 30 nm in diameter. The genome consists of two double-stranded RNA segments (King et al. 2012).

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708

Cucumis sativus (Cucumber)

Cucumber fruit mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CFMMV)

Family: Virgaviridae

Geographical distribution CFMMV infection in plants of Cucumis sativus was reported from Israel (Antignus et al. 2001). Symptoms and host(s) The virus-infected cucumber plants exhibit bright mottling and mosaic on the fruit and severe mosaic vein-banding and yellow mottling on the leaves. Transmission The virus is mechanically sap-transmissible to species belonging to Cucurbitaceae. The virus is transmissible by contact between plants. There is no known vector for this virus. Virion properties and genome The virions are non-enveloped, rigid helical rods with a helical symmetry, about 18 nm in diameter, and 300–310 nm in length. The genome is a monopartite, linear, positive-sense ssRNA of 6562 nt (AF321057 = NC_002633). The 30 terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G50 pppG) (Antignus et al. 2001; Zaitlin 2011; Adams et al. 2017).

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

Geographical distribution CGMMV infection was first reported in plants of Cucumis sativus by Ainsworth (1935) from Great Britain. The virus spreads in the Eurasian region, Korea, France, Greece, Israel, Syria, Spain, Iran, India, Pakistan, Japan, Canada, China, Poland, Taiwan, and the UK (Lovisolo 1981; Avgelis and Vovlas 1986; Hseu et al. 1987; Celix et al. 1996; Varveri et al. 2002; Budzanivska et al. 2006; Yoon et al. 2008; Liu et al. 2009; Moradi and Jafarpour 2011; Ali et al. 2014; Ling et al. 2014; Borodynko-Filas et al. 2017; Crespo et al. 2017; Kumar et al. 2017; Nagendran et al. 2017). Symptoms and host(s) The virus-infected cucumber plants exhibit symptoms of green mottle mosaic and necrotic spots. In the early stages of infection, young leaves of infected cucumber plants displayed light green mottle and blisters. The infected plants were stunted in growth, with darker green blisters and green mottle mosaic symptoms on mature leaves (Komuro 1971; Mandal et al. 2008). CGMMV naturally infects several cucurbits, e.g., Cucumis sativus, Cucumis anguria, Cucumis melo, Citrullus lanatus, and Lagenaria siceraria. In Antarctica, CGMMV has been detected in unusual hosts, mosses (Barbilophozia and Polytrichum) and Antarctic hair grass (Deschampsia antarctica) (Varma and Giri 1998). The experimental host range of CGMMV is narrow and mainly restricted to Cucurbitaceae, Chenopodiaceae, and a few species of Solanaceae. Momordica charantia and N. tabacum are symptomless hosts.

Cucumis sativus (Cucumber)

709

Transmission The virus is not transmitted by any insect vector (Rao and Varma 1984). However Darzi et al. (2018) from Israel have reported that the honeybee Apis mellifera contributes to CGMMV spread via pollination. The virus is transmissible through soil and irrigation water contaminated with infected plant debris (Dorst 1988; Paludan 1985; Vani and Varma 1993). The virus is transmissible by contact between plants. The virus is seed-transmitted in Cucumis sativus to the extent of 4.2% (Kawai et al. 1985; Murthy et al. 2008). The virus is also established to be pollen-transmitted (Liu et al. 2014). Virion properties and genome The virions are non-enveloped, rigid helical rods with a helical symmetry, about 18 nm in diameter, and 300 nm in length. The genome is a monopartite, linear, positive-sense ssRNA of 6424 nt (D12505 = NC_001801). The 30 terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G50 pppG) (Ugaki et al. 1991; Kim et al. 2003; Moradi and Jafarpour 2011; Zaitlin 2011; Li et al. 2016; Adams et al. 2017).

Cucumber leaf spot virus Taxonomic position Genus: Aureusvirus

(CLSV)

Family: Tombusviridae

Geographical distribution CLSV infection was first reported in plants of Cucumis sativus from Germany by Weber et al. (1982). The virus spreads in Bulgaria, Israel, Iran, Spain, France, Germany, Greece, Jordan, Poland, and the UK (Kostova et al. 2001a; Segundo et al. 2001; Pospieszny et al. 2004; Rosner et al. 2006; Bananej et al. 2014a). Symptoms and host(s) The virus was reported to induce numerous chlorotic spots with necrotic centers on the leaves of cucumber plants grown in a commercial greenhouse in southeastern Spain (Segundo et al. 2001). In Poland, CLSV was detected in two commercial greenhouses during 2003, where infected cucumber leaves exhibited sporadic chlorotic spots sometimes with necrotic centers (Pospieszny et al. 2004). The infected young plants were stunted and flowering was delayed. At a later stage, the symptom may disappear. Under experimental conditions, CLSV induces local and systemic infections on Cucumis sativus, Nicotiana benthamiana, and N. clevelandii and induced only local necrotic lesions on Chenopodium quinoa, C. amaranticolor, C. ficifolium, C. murale, Petunia hybrida, Cucurbita melo, Zinnia elegans, and Spinacia oleracea (Pospieszny et al. 2004). Transmission The virus is transmitted by fungal vectors, Olpidium bornovanus and O. radicale (Campbell et al. 1991). The virus is transmissible by mechanical sap-inoculation to three to nine families. The virus is not transmissible by contact between plants. The virus is transmitted by seed. Rosner et al. (2006) have also reported the virus transmission through the recycled drainage water.

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Cucumis sativus (Cucumber)

Virion properties and genome The virions are isometric, non-enveloped, and 30 nm in diameter and have T = 3 icosahedral symmetry. The genome is a monopartite, linear, single-stranded, positive-sense RNA of 4432 nt (DQ227315 = NC_007816) and contains five ORFs; the coat protein contains a P domain (Miller et al. 1997; Martelli et al. 1998; Reade et al. 2003; Martelli and Rubino 2011).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

Geographical distribution CMV infection in plants of Cucumis sativus was first reported in 1916 as the causal agent of plant diseases (Doolittle 1916). The virus is probably distributed worldwide (Magid 1991; Daniels and Campbell 1992; Palukaitis et al. 1992; Eiras et al. 2004; Raj et al. 2008; Zitter and Murphy 2009; Elbeshehy and Sallam 2012; Kumari et al. 2013; Hema et al. 2015; Hossein Vafaei and Mahmoodi 2015). Symptoms and host(s) In virus-infected cucumber plants, the first symptom is a clearing of veins, followed by development of mosaic patterns or mottling consisting of irregularly shaped, dark green areas alternating with light green or yellow areas. The leaves on some species and cultivars are drastically reduced in size, and growth is often retarded. Malformations of the fruit can also occur. The virus has a wide host range and infects vegetables, ornamentals, and other economically important crops, including as many as 1200 host species in 100 families. Two major subgroups of isolates are found (sgI, sgII), with sgI further divided into sgIA and sgIB; isolates of sgIB are more common in Asia (Magid 1991; Palukatis et al. 1992; Eiras et al. 2004; Lee et al. 2008). Transmission The virus is transmitted by aphid vectors: more than 80 species including Myzus persicae, Aphis gossypii, Acyrthosiphon pisum, and Aphis craccivora in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation. Seed transmission in cucumber (1.4%) was reported as early as 1920 by Doolittle, and presently, virus is known to be transmitted through the seeds of at least 19 plant species (Doolittle 1920; Magid 1991). Virion properties and genome The virions are non-enveloped, spherical particles of about 25–30 nm in diameter with T = 3 icosahedral symmetry, and composed of 180 coat proteins: 12 pentamers and 20 hexamers. The genome is segmented, tripartite, linear, positive-sense ssRNA composed of RNA-1, RNA-2, and RNA-3. RNA-1 is 3389 nt (X02733; D00356 = NC_002034), RNA-2 is 3050 nt (D00355 = NC_002035), and RNA-3 2197 nt (M21464; D10538 = NC_001440). A sub-genomic RNA-4 is also encapsidated in the particles that contain RNA-3. The coat protein consists of 218 amino acids with a molecular weight of 24.2 kDa (Ding et al. 1994; Palukaitis and Garcia-Arenal 2003). For RNAs 1, 2, and 3, there is about 78%, 74%, and 78% identity between CMV subgroups, respectively (Gould and Symons 1982; Rezaian et al. 1985; Rizzo and Palukaitis 1988, 1989; Garcia-Arenal Rodriguez and Fraile 2011; Scholthof et al. 2011; Scott 2011; Elbeshehy and Sallam 2012).

Cucumis sativus (Cucumber)

Cucumber mottle virus Taxonomic position Genus: Tobamovirus

711

(CuMoV)

Family: Virgaviridae

Geographical distribution CuMoV infection in plants of Cucumis sativus was reported from Japan (Orita et al. 2006). Symptoms and host(s) The virus-infected cucumber plants exhibit mottling and vein-clearing symptoms. Transmission The virus is mechanically sap-transmissible. The virus is also transmissible by contact between plants. Virion properties and genome The virions are non-enveloped, rigid helical rods with a helical symmetry, about 18 nm in diameter and 300–310 nm in length. The genome is a monopartite, linear, positive-sense ssRNA of 6485 nt (AB261167 = NC_008614). The 30 terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G50 pppG) (Orita et al. 2006; Zaitlin 2011; Adams et al. 2017).

Cucumber necrosis virus Taxonomic position Genus: Tombusvirus

(CNV)

Family: Tombusviridae

Geographical distribution CNV infection in plants of Cucumis sativus was first reported from Ontario, Canada, by McKeen (1959). The virus spreads in Canada (Ontario) (Rochon and Tremaine 1989; Li et al. 2013). Symptoms and host(s) The virus-infected cucumber plants exhibit chlorosis, necrosis, and malformation in the leaves. Transmission The virus is transmitted by the soil-borne fungi, Olpidium bornovanus and Olpidium cucurbitacearum, through binding of the virus on the surface of zoospores of the fungus (Dias 1970; Li et al. 2013). The virus is transmissible by mechanical sap-inoculation, and also through contact between plants, but not transmitted by seed or pollen. Virion properties and genome The virions are isometric and icosahedral (T = 3), non-enveloped, and about 32–35 nm in diameter. The genome is a monopartite, linear, single-stranded, positive-sense RNA of 4701 nt containing five ORFs (M25270 = NC_001469) (Rochon and Tremaine 1988, 1989; White 2011).

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Cucumis sativus (Cucumber)

Cucumber soil-borne virus Taxonomic position Genus: Unassigned

(CuSBV)

Family: Tombusviridae

Geographical distribution CuSBV infection in plants of Cucumis sativus was reported from Jieh, south of Beirut, Lebanon, by Koenig et al. (1983). Symptoms and host(s) The virus-infected cucumber plants do not exhibit visual systemic symptoms on leaves, but the virus is present in roots only. Transmission The virus is transmitted by a fungal vector, Olpidium bornovanus. The virus is also transmissible by mechanical sap-inoculation to less than three plant families. Virion properties and genome The virions are isometric and icosahedral (T = 3), non-enveloped, and 28–34 nm in diameter. The genome is a monopartite, linear, single-stranded, positive-sense RNA. No sequence information is currently available.

Cucumber vein-clearing virus Taxonomic position Genus: Carlavirus

(CuVCV)

Family: Betaflexiviridae

Geographical distribution CuVCV infection in plants of Cucumis sativus was reported from Arusha and Tanzania (Menzel et al. 2011). Symptoms and host(s) The virus-infected cucumber plants exhibit vein-clearing symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci B. biotype, in a semi-persistent manner (Menzel et al. 2011). The virus is also mechanically sap-transmissible. Virion properties and genome The virions are slightly flexuous filaments, 610–700 nm in length, and 12–15 nm in diameter. The genome consists of a single molecule of positive-sense ssRNA. It has a partial genome sequence of 5218 nt (JN591720), excluding the 30 poly(A) tail is available (Adams et al. 2004).

Cucumis sativus (Cucumber)

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Cucumber vein yellowing virus Taxonomic position Genus: Ipomovirus

(CVYV)

Family: Potyviridae

Geographical distribution CVYV was first described in Israel in the late 1950s, causing a severe cucumber disease in the Jordan Valley (Cohen and Nitzany 1960). The virus infection in plants of Cucumis sativus was reported from Iran, Israel, Jordan, Greece, Portugal, Spain, Sudan, Syria, Saudi Arabia, and Turkey (Al-Musa et al. 1985; Yilmaz et al. 1989; Cuadrado et al. 2001; Louro et al. 2004; CABI/EPPO 2005; Bananej et al. 2006; Fidan et al. 2012; Abrahamian et al. 2013; Bananej et al. 2014b). Symptoms and host(s) The virus-infected cucumber plants exhibit symptoms of pronounced vein-clearing, mottling, and deformation of leaves followed by generalized chlorosis and necrosis of affected tissues and finally leading to death of plants. Fruits show yellow/green chlorotic spots and sometimes light to dark green mottling. The natural host range of CVYV was first thought to be limited to cucurbits. Recent studies showed that CVYV could also infect several non-cucurbit weeds under natural conditions including the perennial Convolvulus arvensis and the annual Malva parviflora (Janssen et al. 2002). Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a non-persistent manner (Harpaz and Cohen 1965; Janssen et al. 2005; Navas-Castillo et al. 2014). The virus is also mechanically saptransmissible (Mansour and Al-Musa 1993). Virion properties and genome The virions are filaments, non-enveloped, and flexuous measuring 740–800 nm in length and 15–18 nm in width. The genome consists of a single molecule of positive-sense ssRNA of 9751 nt (AY578085 = NC_006941) (Mansour and Hadidi 1999; Lecoq et al. 2000; Janssen et al. 2005; Colinet 2011; Galipienso et al. 2012; Wylie et al. 2017).

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(CABYV)

Family: Luteoviridae

CABYV infection in plants of Cucumis sativus was reported from the Czech Republic, France, Greece, Iran, Spain, Egypt, Tunisia, Turkey, Korea, Saudi Arabia, India, and Lebanon (Lecoq et al. 1992; AbouJawdah et al. 1997; Lecoq 1999; Hattab et al. 2005; Boubourakas et al. 2006; Yardimci and Ozgonen 2007; Xiang et al. 2008; Hattab et al. 2009; Lecoq and Desbiez 2012; Salehi et al. 2012; Al-Saleh et al. 2015b; Bananej et al. 2015; Choi et al. 2015; Suveditha et al. 2017). The virus-infected cucumber plants show yellowing and thickening of the older leaves; symptom intensity differs depending upon cultivar and season. The virus is transmitted by aphid vectors, Myzus persicae, M. euphorbiae, and Aphis gossypii, in a persistent, circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation (Lecoq et al. 1992). For more details of CABYV, refer to Cucumis melo.

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Cucurbit chlorotic yellows virus

(CCYV)

Taxonomic position CCYV is a tentative member of the genus Crinivirus and family Closteroviridae CCYV infection in plants of Cucumis sativus was reported from China, Taiwan, Greece, Japan, Sudan, Lebanon, Saudi Arabia, Turkey, and Iran (Huang et al. 2010; Okuda et al. 2010; Gu et al. 2011; Hamed et al. 2011; Abrahamian et al. 2012, 2015; Bananej et al. 2013; Orfanidou et al. 2014; Al-Saleh et al. 2015a; Orfanidou et al. 2017). The virus-infected cucumber plants exhibit symptoms of chlorosis, yellowing, and interveinal chlorotic spots on lower leaves. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a semi-persistent manner (Furuta et al. 2008). For more details of CCYV, refer to Cucumis melo.

Cucurbit yellow stunting disorder virus Taxonomic position Genus: Crinivirus

(CYSDV)

Family: Closteroviridae

CYSDV infection in plants of Cucumis sativus was reported from Israel, Jordan, Morocco, Portugal, Spain, Turkey, Greece, France, Lebanon, Saudi Arabia, Japan, Syria, UAE, Egypt, Mexico, Saudi Arabia, and the USA (Livieratos et al. 1999; Rubio et al. 1999; Desbiez et al. 2000, 2003; Louro et al. 2000; Papayiannis et al. 2005; Boubourakas et al. 2006; Fidan et al. 2012; Abrahamian et al. 2015). The virus-infected cucumber plants show severe chlorotic mottling and yellowing symptoms that start as an interveinal mottle on the older leaves and intensifying as leaves mature: infected plants are also stunted (Abou-Jawdah et al. 2000). The virus is transmitted by the whitefly vector Bemisia tabaci in a semipersistent manner. The virus is not mechanically sap-transmissible. For more details of CYSDV, refer to Cucumis melo.

Eggplant mottled dwarf nucleorhabdovirus

(EMDV)

Synonyms Cucumber toad skin virus Taxonomic position Genus: Nucleorhabdovirus

Family: Rhabdoviridae

EMDV was reported in plants of Cucumis sativus early in the 1980s in Southeastern France by Lecoq (1982). The virus spreads in Greece, Italy, Bulgaria, and Spain (Roggero et al. 1995; Katis et al. 2000; Kostova et al. 2001b; Aramburu et al. 2006). The virus-infected cucumber plants show vein-clearing on older leaves and a severe inward curling of younger leaves with the main vein showing a sinuous course giving a typical “toad skin”-like appearance. The infected plants are stunted with shortened internodes. The virus is transmitted by leafhopper vectors, Anaceratagallia laevis and A. ribauti, in a persistentpropagative manner (Della Giustina et al. 2000). The virus is mechanically sap-transmissible to solanaceous hosts. For more details of EMDV, refer to Solanum melongena.

Cucumis sativus (Cucumber)

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Groundnut ringspot orthotospovirus Taxonomic position Genus: Orthotospovirus

(GRSV)

Family: Tospoviridae

GRSV infection in plants of Cucumis sativus was reported from Brazil (Spadotti et al. 2014). The virusinfected cucumber plants exhibit symptoms of necrotic concentric rings on leaves and fruits. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sapinoculation. For more details of GRSV, refer to Arachis hypogaea.

Hop stunt viroid

(HpSVd)

Synonyms Cucumber pale fruit viroid Taxonomic position Genus: Hostuviroid

Family: Pospiviroidae

Pale fruit disease of Cucumis sativus plants was first reported from the Netherlands in 1963 (Van Dorst and Peters 1974). The viroid spreads in the Netherlands and Finland (Lemmetty et al. 2011; Yaguchi and Takahashi 1984; Martinez et al. 2010). The viroid-infected cucumber plants exhibit stunting of the whole plant, leaf curling, flower deformation, and pale-colored fruits. The viroid is easily transmissible mechanically from cucumber to cucumber by sap-inoculation, but is not transmissible through seed, soil, and aphids (Van Dorst and Peters 1974; Sano et al. 1984; Puchta et al. 1988). For more details of HpSVd, refer to Humulus lupulus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in greenhouse-grown plants of Cucumis sativus was reported from Italy (Vicchi et al. 1999). The virus is transmitted mainly by a thrips vector, Frankliniella occidentalis, in a persistentpropagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Kyuri green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(KGMMV)

Family: Virgaviridae

Geographical distribution KGMMV infection was first reported in Cucumis sativus from Japan by Inouye et al. (1967). The virus spreads in Japan, Korea, and Indonesia (Francki et al. 1986; Tan et al. 2000).

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Symptoms and host(s) The virus-infected cucumber plants exhibit symptoms of mottling mosaic, severe fruit mottling, and malformation. Transmission The virus is transmitted by means not involving a vector. The virus is transmissible by mechanical sapinoculation to more than nine plant families. The virus is transmitted through seeds and contaminated soil. The virus is also transmissible by contact between plants. Virion properties and genome The virions are rod-shaped, non-enveloped, and usually straight with a clear modal length of c. 300 nm and c. 15 nm wide (Tan et al. 2000). The genome consists of a single molecule of positive-sense ssRNA of 6514 nt (AJ295948 = NC_003610) and contains four open reading frames (ORFs) encoding 131, 189, 28, and 18 kDa proteins (Yoon et al. 2001). The 30 terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G50 pppG) (Zaitlin 2011; Adams et al. 2017).

Melon necrotic spot virus Taxonomic position Genus: Gammacarmovirus

(MNSV)

Family: Tombusviridae

MNSV infection in plants of Cucumis sativus was reported from Northern England, the Netherlands, Crete, and Iran (Bos et al. 1984; Tomlinson and Thomas 1986; Avgelis 1990; Safaeezadeh 2007). The virus-infected cucumber plants exhibit vein-clearing and chlorotic leaf spotting which develops into desiccating necrotic spots. The virus is transmitted by the fungal vector Olpidium bornovanus. The virus is transmissible by mechanical sap-inoculation, and also by contact between plants. For more details of MNSV, refer to Cucumis melo.

Melon yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(MYSV)

Family: Tospoviridae

MYSV infection in plants of Cucumis sativus was reported from Ecuador, Japan, and Taiwan (Takeuchi et al. 2001; Okuda et al. 2002; Chao et al. 2010; Quito-Avila et al. 2014). The virus-infected cucumber plants develop yellow mosaic on the upper leaves and yellow spots on the lower leaves, which coalesce; later necrosis develops. Severely affected plants become stunted. The virus is transmitted by a thrips vector, Thrips palmi, in a persistent-propagative manner, and also through mechanical sap-inoculation (Sugiyama et al. 2009). For more details of MYSV, refer to Cucumis melo.

Mungbean yellow mosaic India virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

(MYMIV)

Cucumis sativus (Cucumber)

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MYMIV infection in plants of Cucumis sativus was reported from Oman (Shahid et al. 2018). The virus-infected cucumber plants exhibit foliar yellow mosaic and crumpling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by mechanical inoculation. For more details of MYMIV, refer to Vigna radiata.

Papaya ringspot virus Taxonomic position Genus: Potyvirus

(PRSV)

Family: Potyviridae

PRSV pathotype W infection in plants of Cucumis sativus was reported from India, Nigeria and the Cook Islands (Owolabi et al. 2008; Nagendran et al. 2017). The virus is transmitted by aphid vectors in a non-persistent manner, and also through mechanical sap-inoculation. For more details of PRSV, refer to Carica papaya.

Potato virus Y

(PVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

PVY infection in plants of Cucumis sativus was reported from Turkey (Ozaslan et al. 2006). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts and by grafting. For more details of PVY, refer Solanum tuberosum.

Squash leaf curl virus Taxonomic position Genus: Begomovirus

(SLCuV)

Family: Geminiviridae

SLCuV infection in plants of Cucumis sativus was reported from Palestine and Lebanon (Ali-Shtayeh et al. 2010; Sobh et al. 2012; Abrahamian et al. 2015). The virus-infected cucumber plants exhibit symptoms of leaf curling at the growing tips which often disappear as the plants mature and begin to flower and set fruit. Symptoms on fruit are not particularly notable, perhaps because very little fruit is set on most cultivars after infection occurs in susceptible cultivars (Ali-Shtayeh et al. 2010). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of SLCuV, refer to Cucurbita pepo.

Squash mosaic virus Taxonomic position Genus: Comovirus

(SqMV)

Family: Secoviridae

SqMV infection in plants of Cucumis sativus was reported from Indonesia (Lestari and Nurhayati 2014). The virus is transmitted by beetle vectors in a non-persistent manner, and also by mechanical

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sap-inoculation. The virus is seed-transmitted in Cucumis sativus up to 100% (Lestari and Nurhayati 2014; Purba et al. 2017). For more details of SqMV, refer to Cucurbita pepo.

Tobacco necrosis virus A Taxonomic position Genus: Alphanecrovirus

(TNV-A)

Family: Tombusviridae

TNV-A infection in plants of Cucumis sativus was reported from Lithuania (Zitikaite and Staniulis 2005). The virus-infected cucumber plants exhibit symptoms of various spotting or mottling symptoms on leaves or fruits and necrotic symptoms on older leaves. The virus is transmitted by a fungal vector, Olpidium brassicae, and also by mechanical sap-inoculation. For more details of TNV-A, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Cucumis sativus is widespread primarily in the USA and Europe (Sirca et al. 2007). The virus-infected cucumber plants exhibit very bright yellow mosaic and ringspot symptoms. Under natural conditions this virus infects a large number of hosts. The virus is transmitted by Xiphinema rivesi in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. The virus is transmitted through the seed of Cucumis sativus. For more details of TRSV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Cucumis sativus was reported from India (Krishnareddy et al. 2003; Pavithra et al. 2016; Suveditha et al. 2017). The virus-infected cucumber plants exhibit symptoms of necrotic lesions on the leaves, extending to mid veins, petioles, flower buds, and tip. This is followed by dieback of veins. The virus is transmitted by the thrips vectors. The virus present in/on pollen enters in to the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato black ring virus Taxonomic position Genus: Nepovirus

(TBRV)

Family: Secoviridae

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TBRV infection in plants of Cucumis sativus was reported from Poland (Pospieszny et al. 2003a). The virus-infected cucumber plants exhibit systemic mottling and necrosis symptoms. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TBRV, refer to Solanum lycopersicum.

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Cucumis sativus was reported from India, Tunisia, Thailand, and Indonesia (Ito et al. 2008; Mizutani et al. 2011; Manjunath 2014; Mnari-Hattab et al. 2015; Suveditha et al. 2017). The virus-infected cucumber plants exhibit severe yellowing/green mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner, and also by grafting. For more details of ToLCNDV, refer to Solanum lycopersicum.

Tomato leaf curl Palampur virus Taxonomic position Genus: Begomovirus

(ToLCPalV)

Family: Geminiviridae

ToLCPalV infection in plants of Cucumis sativus was reported from Iran (Heydarnejad et al. 2009, 2013). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner, and also by grafting. For more details of ToLCPalV, refer to Solanum lycopersicum.

Tomato mosaic virus Taxonomic position Genus: Tobamovirus

(ToMV)

Family: Virgaviridae

ToMV infection in plants of Cucumis sativus was reported from Iran (Yazdani-Khameneh et al. 2013). No vector is involved in the spread of this virus. This virus is transmissible by mechanical sapinoculation, transmitted by grafting, and also by contact between plants. For more details of ToMV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Cucumis sativus was reported from Europe (Sirca et al. 2007). The virusinfected cucumber plants exhibit diffuse green yellow mosaic with some rugosity. The plants gradually recover from symptoms. The virus is transmitted by a nematode vector, Xiphinema rivesi, in a

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non-persistent manner, and also by mechanical sap-inoculation. For more details of ToRSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Cucumis sativus has been reported from Bulgaria, Israel, and Zimbabwe (Gera et al. 2000; Karadjova et al. 2003; Karavina and Gubba 2017). The virus-infected cucumber plants exhibit chlorotic local lesions on the leaves and leaf deformation symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Cucumis sativus has been reported from Kuwait (Al-Ali et al. 2016). The virus-infected cucumber plants exhibit upward leaf curling, chlorosis, plant dwarfing, mosaic, and yellowing, as well as fruit and leaf deformation. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

Watermelon chlorotic stunt virus Taxonomic position Genus: Begomovirus

(WmCSV)

Family: Geminiviridae

WmCSV infection in plants of Cucumis sativus was reported from Lebanon (Samsatly et al. 2012). The virus-infected cucumber plants exhibit yellowing symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by mechanical inoculation. The virus is transmissible by grafting. For more details of WmCSV, refer to Citrullus lanatus.

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV-1 isolate infection in plants of Cucumis sativus was reported from India, Iran, Taiwan, and Lebanon (Makkouk and Lesemann 1980; Thomas 1980; Dikova et al. 1984; Hseu et al. 1987; Hossein

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Vafaei and Mahmoodi 2015; Kumar et al. 2015). The virus-infected cucumber plants exhibit severe mottling, blistering, and malformation symptoms. The virus is transmitted by the green peach aphid, Myzus persicae, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of WMV, refer to Citrullus lanatus.

Watermelon silver mottle orthotospovirus Taxonomic position Genus: Orthotospovirus

(WSMoV)

Family: Tospoviridae

WSMoV infection in plants of Cucumis sativus was reported from Taiwan and Japan (Okuda et al. 2002). The virus-infected cucumber plants exhibit symptoms of silver mottling on leaves and chlorotic mottling on fruits. The virus is transmitted by the thrips vector, Thrips palmi, in a persistent-propagative manner, and also by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants. For more details of WSMoV, refer to Citrullus lanatus.

Zucchini lethal chlorosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(ZLCV)

Family: Tospoviridae

ZLCV infection in plants of Cucumis sativus was reported from Brazil (Nagata et al. 1998). The virusinfected cucumber plants exhibit symptoms of yellowing, mottling, and vein-banding on the leaves. The virus is transmitted by a thrips vector, Frankliniella zucchini, in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of ZLCV, refer to Cucurbita pepo.

Zucchini tigre mosaic virus Taxonomic position Genus: Potyvirus

(ZTMV)

Family: Potyviridae

ZTMV infection in plants of Cucumis sativus was reported from China (Xiao et al. 2016). The virusinfected cucumber plants exhibit mosaic mottling symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ZTMV refer to Cucurbita pepo.

Zucchini yellow fleck virus Taxonomic position Genus: Potyvirus

(ZYFV)

Family: Potyviridae

ZYFV infection in plants of Cucumis sativus was reported from Greece, Iran, and certain Mediterranean countries (Avgelis 1985; Gilbert-Albertini et al. 1995; Safaeezadeh 2007). The virus-infected cucumber plants exhibit severe mosaic symptoms. The virus is transmitted by an aphid vector,

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Cucumis sativus (Cucumber)

Myzus persicae, in a non-persistent manner, and also through mechanical sap-inoculation. For more details of ZYFV, refer to Cucurbita pepo.

Zucchini yellow mosaic virus Taxonomic position Genus: Potyvirus

(ZYMV)

Family: Potyviridae

ZYMV infection in plants of Cucumis sativus was reported from India, Iran, Saudi Arabia, Lebanon, Ontario, Singapore, Poland, Germany, Turkey and the Czech Republic (Lesemann et al. 1983; Hseu et al. 1985, 1987; Stobbs et al. 1990; Wong et al. 1994; Al-Shahwan et al. 1995; Desbiez and Lecoq 1997; Svoboda and Polak 2002; Pospieszny et al. 2003b; Verma et al. 2004; Muller et al. 2006; Ozaslan et al. 2006; Hossein Vafaei and Mahmoodi 2015; Karthikeyan et al. 2017). The virus-infected cucumber plants show vein-clearing, yellow mosaic, yellowing, shoestring leaves, and stunting. The infected young cucumber fruits were curved and hooked. The virus is transmitted by many Aphid spp. in a nonpersistent manner (Katis et al. 2006). The virus is also mechanically sap-transmissible and infects mostly cultivated and wild cucurbits, few ornamentals, and weeds. For more details of ZYMV, refer to Cucurbita pepo.

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Rochon DM, Tremaine JH (1989) Complete nucleotide sequence of the cucumber necrosis virus genome. Virology 169:251–259 Roggero P, Milne RG, Masenga V, Ogliara P, Stravato VM (1995) First reports of Eggplant mottled dwarf rhabdovirus in cucumber and in pepper. Plant Dis 79(3):321 Rosner A, Lachman O, Pearlsman M, Feigelson L, Maslenin L, Antignus Y (2006) Characterisation of cucumber leaf spot virus isolated from recycled irrigation water of soil-less cucumber cultures. Ann Appl Biol 149:313–316 Rubio L, Soong J, Kao J, Falk BW (1999) Geographic distribution and molecular variation of isolates of 3 whitefly-borne closteroviruses: of cucurbits, Lettuce infectious yellows virus, Cucurbit yellow stunting disorder virus and Beet pseudo yellows virus. Phytopathology 89:707–711 Safaeezadeh M (2007) First report of Melon necrotic spot virus and Zucchini yellow fleck virus in cucurbits in Iran. J Plant Pathol 89:302 Salehi S, Vahdat A, Bananej K, (2012) Occurrence of Cucurbit aphid-borne yellows virus associated with yellowing symptoms on greenhouse-grown cucumber in Alborz and Tehran provinces. Proceedings 20th Iranian Plant Protection Congress, Shiraz, Iran, 856 Samsatly J, Sobh H, Jawhari M, Najjar C, Haidar A, Abou-Jawdah Y (2012) First report of watermelon chlorotic stunt virus in cucurbits in Lebanon. Plant Dis 96:1703 Sano T, Uyeda I, Shikata E, Ohno T, Okada Y (1984) Nucleotide sequence of cucumber pale fruit viroid: homology to hop stunt viroid. Nucleic Acids Res 12:3427–3434 Scholthof K-BG, Adkins S, Czosnek H, Palukaitis P, Jacquot E, Hohn T, Hohn B, Saunders K, Candresse T, Ahlquist P, Hemenway C, Foster AGD (2011) Top 10 plant viruses in molecular plant pathology. Mol Plant Pathol 12:938–954 Scott SW (2011) Bromoviridae and allies. In: Encyclopedia of Life Sciences (ELS). Wiley, Chichester. https://doi.org/ 10.1002/9780470015902 Segundo E, Janssen D, Velasco L, Ruiz L, Cuadrado IM (2001) First report of cucumber leaf spot virus in Spain. Plant Dis 85:1123 Shahid MS, Al-Mahmooli IH, Al-Sadi AM, Briddon RW (2018) Identification of Mungbean yellow mosaic India virus infecting cucumber in Oman. Plant Dis 102:465 Sirca S, Geric Stare B, Mavric Plesko I, Virscek Marn M, Urek G, Javornik B (2007) Xiphinema rivesi from Slovania transmit Tobacco ringspot virus and Tomato ringspot virus to cucumber bait plants. Plant Dis 91:770 Sobh H, Samsatly J, Jawhari M, Najjar C, Haidar A, Abou-Jawdah Y (2012) First report of Squash leaf curl virus in cucurbits in Lebanon. Plant Dis 96(8):1231 Spadotti DMA, Leao EU, Rocha KCG, Pavan MA, Krause-Sakate R (2014) First report of groundnut ringspot virus in cucumber fruits in Brazil. New Dis Rep 29:25 Stobbs LW, van Schagen JG, Shantz GM (1990) First report of zucchini yellow mosaic virus in Ontario. Plant Dis 74:394 Sugiyama M, Okuda M, Sakata Y (2009) Evaluation of resistance to melon yellow spot virus in a cucumber germplasm collection. Plant Breed 128:696–700 Suveditha S, Swarnalatha V, Krishna Reddy M (2017) Identification and molecular diagnosis of viruses infecting cucumber. In: 26 annual Conference of Indian Virological Society (VIROCON 2017), Held at Mangaluru, India. p 124 Svoboda J, Polak J (2002) Distribution, variability and over wintering of Zucchini yellow mosaic virus in the Czech Republic. Plant Prot Sci 38:125–130 Takeuchi S, Okuda M, Hanada K, Kawada K, Kameya-Iwaki M (2001) Spotted wilt disease of cucumber (Cucumis sativus) caused by Melon yellow spot virus. Jpn J Phytopathol 67:46–51 Tan SH, Nishiguchi M, Murata M, Motoyoshi F (2000) The genome structure of kyuri green mottle mosaic tobamovirus and its comparison with that of cucumber green mottle mosaic tobamovirus. Arch Virol 145:1067–1079 Thomas W (1980) Watermelon mosaic virus, the cause of a serious disease of cucurbits in Cook Islands. N Z J Exp Agric 8:309–312 Tomlinson JA, Thomas BJ (1986) Studies on melon necrotic spot virus disease of cucumber and on the control of the fungus vector (Olpidium radicale). Ann Appl Biol 108:71–80 Ugaki M, Toomiyama M, Kakutani T, Hidaka S, Kiguchi T, Nagata R, Sato T, Motoyoshi F, Nishiguchi M (1991) The complete nucleotide sequence of cucumber green mottle mosaic virus (SH strain) genomic RNA. J Gen Virol 72:1487–1495 Van Dorst HJM, Peters D (1974) Some biological observations on pale fruit, a viroid-incited disease of cucumber. Neth J Plant Pathol 80:85–96 Vani S, Varma A (1993) Properties of Cucumber green mottle mosaic virus isolated from water of river Jamuna. Indian Phytopathol 46:118–122 Varma A, Giri BK (1998) Virus diseases of cucurbits. In: Nayar NM, More TA (eds) Cucurbit. Oxford and IBH Publishing House Private Ltd., New Delhi, pp 225–245 Varveri C, Vassilakos N, Bem F (2002) Characterization and detection of cucumber green mottle mosaic virus in Greece. Phytoparasitica 30:493–501 Verma R, Prakash S, Tomer SPS (2004) First report of Zucchini yellow mosaic virus in cucumber (Cucumis sativus) in India. Plant Dis 88:906

Cucurbita foetidissima (Buffalo gourd)

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Vicchi V, Fini P, Cardoni M (1999) Presence of Impatiens necrotic spot tospovirus (INSV) on vegetable crops in EmiliaRomagna region. Informatore Fitopatol 49(4):53–55 Weber I, Proll E, Ostermann W-D, Leiser R-M, Stanarius A, Kegler H (1982) Description of Cucumber leaf spot virus, a virus unknown so far in greenhouse cucumber. Arch Phytopathol Pflanzenschutz 18:137–154 White KA (2011) Tombusvirus. Tombusviridae. In: The Springer index of Viruses. Springer, New York, pp 1917–1926. https://doi.org/10.1007/978-0-387-95919-1_314 Wisler GC, Dufus JE, Liu H-Y, Li RH (1998) Ecology and epidemiology of whitefly-transmitted closteroviruses. Plant Dis 82:270–280 Wong SM, Chung CG, Chung CY, Chong PL (1994) Characterization of an isolate of Zucchini yellow mosaic virus from cucumber in Singapore. J Phytopathol 141:335–368 Woudt LP, de Rover AP, de Haan PT, van Grinsven MQJM (1993) Sequence analysis of the RNA genome of cucumber chlorotic spot virus (CCSV), a whitefly transmitted closterovirus. In: Abstracts, IX Int. Congress of Virology, Glasgow, pp 326 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J GenVirol 98:352–354 Xiang HY, Shang QX, Han CG, Li DW, Yu JL (2008) First report on the occurrence of Cucurbit aphid-borne yellows virus on nine cucurbitaceous species in China. Plant Pathol 57:390 Xiao L, Li YY, Tan GL, Lan PX, Zhong L, Liu Y, Li R, Li F (2016) First report of Zucchini tigre mosaic virus infecting several cucurbit plants in China. Plant Dis 100:1253 Yaguchi S, Takahashi T (1984) Response of cucumber cultivars and other Cucurbitaceous species to infection by Hop stunt viroid. Phytopathol Z 109:21–32 Yamashita S, Doi Y, Yora K, Yoshino M (1979) Cucumber yellows virus: its transmission by the greenhouse whitefly, Trialeurodes vaporariorum (Westwood), and the yellowing disease of cucumber and muskmelon caused by the virus. Ann Phytopathol Soc Jpn 45:484–496 Yardimci N, Ozgonen H (2007) First report of cucurbit aphid-borne yellows virus in Turkey. Aust Plant Dis Notes 2:59 Yazdani-Khameneh S, Hamedi A, Farahani AA, Hashemi S, Golnaraghi AR, Rakhshandehroo F (2013) Tomato mosaic virus on cucumber and potato in Iran. J Plant Pathol 95:659–668 Yilmaz MA, Ozaslan M, Ozaslan D (1989) Cucumber vein yellowing virus in cucurbitaceae in Turkey. Plant Dis 73:610 Yoon JY, Min BE, Choi SH, Ryu KH (2001) Completion of nucleotide sequence and generation of highly infectious transcripts to cucurbits from full-length cDNA clone of Kyuri green mottle mosaic virus. Arch Virol 146(11):2085–2096 Yoon JY, Choi GS, Choi SK, Hong JS, Choi JK, Kim W, Lee GP, Ryu KH (2008) Molecular and biological diversities of cucumber green mottle mosaic virus from cucurbitaceous crops in Korea. J Phytopathol 156:408–412 Zaitlin M (2011) Tobamovirus. In: Tidona C, Darai G (eds) The Springer index of viruses. Springer, New York. https://doi. org/10.1007/978-0-387-95919-1 Zenbayashi R, Shimazaki Y, Shibukawa S (1984) Studies on cucumber yellows disease. Bull Saitama Hortic Exp Stn 13:11–40 Zitikaite I, Staniulis J (2005) Isolation and characterization of Tobacco necrosis virus detected on some vegetable species. Biologia 55:35–39 Zitter TA, Murphy JF (2009) Cucumber mosaic. Plant Health Instr. https://doi.org/10.1094/PHI-I-2009-0518-01

Cucurbita foetidissima (Buffalo gourd) Family: Cucurbitaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Commercial crop

(CMV)

Family: Bromoviridae

CMV infection in plants of Cucurbita foetidissima was reported from Arizona (USA) (Rosemeyer 1981, 1982; Rosemeyer et al. 1986). The virus-infected buffalo gourd plants exhibit severe systemic mosaic symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner,

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and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Lettuce infectious yellows virus Taxonomic position Genus: Crinivirus

(LIYV)

Family: Closteroviridae

LIYV infection in plants of Cucurbita foetidissima was reported from Arizona (USA) (Rosemeyer et al. 1982, 1986). The virus-infected buffalo gourd plants exhibit symptoms of slight stunting and extremely mild chlorosis of older leaves. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a semipersistent manner. The virus is not mechanically sap-transmissible. For more details of LIYV, refer to Lactuca sativa.

Squash mosaic virus Taxonomic position Genus: Comovirus

(SqMV)

Family: Secoviridae

SqMV infection in plants of Cucurbita foetidissima was reported from Arizona (USA) (Rosemeyer 1981, 1982; Rosemeyer et al. 1982, 1986). The virus-infected buffalo gourd plants exhibit mild systemic mosaic and vein-clearing symptoms. The virus is transmitted by a number of beetle vectors and also by mechanical sap-inoculation. For more details of SqMV, refer to Cucurbita pepo.

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV infection in plants of Cucurbita foetidissima was reported from Arizona (USA) (Rosemeyer et al. 1982, 1986). The virus-infected buffalo gourd plants exhibit mosaic symptoms on the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of WMV, refer to Citrullus lanatus.

References Rosemeyer ME (1981) Virus studies with Cucurbita foetidissima. Cucurbit Genet Rep 4:41–42 Rosemeyer ME (1982) Isolation and characterization of two viruses from Cucurbita foetidissma HBK, buffalo gourd. MS thesis, University of Arizona, Tucson. p 77 Rosemeyer ME, Wells BH, Zaid A (1982) Diseases of buffalo gourd, Cucurbita foetidissma, in Arizona. Phytopathology 72:955 Rosemeyer ME, Brown JK, Nelson MR (1986) Five viruses isolated from field grown buffalo gourd (Cucurbita foetidissima), a potential crop for semi arid lands. Plant Dis 70:405–409

Cucurbita maxima (Pumpkin)

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Cucurbita maxima (Pumpkin) Family: Cucurbitaceae

Vegetable

Beet pseudoyellows virus Taxonomic position Genus: Crinivirus

C

(BPYV)

Family: Closteroviridae

BPYV infection in plants of Cucurbita maxima was reported from California, USA (Wintermantel 2004). The virus-infected pumpkin plants exhibit extensive leaf chlorosis symptoms. The virus is transmitted by the whitefly vector, Trialeurodes vaporariorum, in a semi-persistent manner. The virus is not transmissible by mechanical sap-inoculation, and also not by contact between plants. For more details of BPYV, refer to Beta vulgaris.

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

CGMMV infection in plants of Cucurbita maxima was reported from India and Pakistan (Ali et al. 2014; Kumar et al. 2017; Nagendran et al. 2017). The virus is not transmitted by any insect vector. The virus is transmissible through soil and irrigation water contaminated with infected plant debris. The virus is pollen transmitted and also by contact between plants. For more details of CGMMV, refer to Cucumis sativus.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Cucurbita maxima was reported from India, Ukraine, and Saudi Arabia (Singh et al. 1991; Zitikaite et al. 2011; Al-Shahwan et al. 2017). The affected pumpkin plants show light green mottled and crinkly foliage. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is seed-transmitted in Cucurbita maxima (Mukhopadhayay and Saha 1968). For more details of CMV, refer to Cucumis sativus.

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

Family: Luteoviridae

(CABYV)

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CABYV infection in plants of Cucurbita maxima was reported from Turkey, Tunisia, and Taiwan (Yardimci and Ozgonen 2007; Hattab et al. 2009; Knierim et al. 2010). The virus-infected pumpkin plants exhibit yellowing symptoms. The virus is transmitted by aphid vectors, Myzus persicae, M. euphorbiae, and Aphis gossypii, in a circulative, non-propagative manner (Lecoq et al. 1992). For more details of CABYV, refer to Cucumis melo.

Cucurbit chlorotic yellows virus

(CCYV)

Taxonomic position CCYV is a tentative member of the genus Crinivirus and family Closteroviridae CCYV infection in plants of Cucurbita maxima was reported from Sudan and Taiwan (Huang et al. 2010; Mohammed et al. 2014). The virus-infected pumpkin plants exhibit yellowing symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a semi-persistent manner. For more details of CCYV, refer to Cucumis melo.

Cucurbit leaf crumple virus

(CuLCrV)

Synonyms Cucurbit leaf curl virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

CuLCrV infection in plants of Cucurbita maxima was reported from Arizona and Texas (USA), and from Mexico (Brown et al. 2000). The virus-infected pumpkin plants exhibit leaf curl and/or mottling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative nonpropagative manner. The virus is not mechanically sap-transmissible. For more details of CuLCrV, refer to Cucumis melo.

Cucurbit vein banding virus Taxonomic position Genus: Potyvirus

(CVBV)

Family: Potyviridae

Geographical distribution CVBV infection in plants of Cucurbita maxima was reported from Argentina (Perotto et al. 2018). Symptoms and host(s) Strong vein banding was observed in leaves of C. maxima in which no other virus was detected (Perotto et al. 2018).

Cucurbita maxima (Pumpkin)

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Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is transmitted by mechanical sap inoculation. The virus is not transmitted by contact between plants; not transmitted by seed; not transmitted by pollen.

Virion properties and genome The virions are nonenveloped, flexuous filaments 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 9969 nt (KY657266 = NC_035134) (Revers and Garcia 2015; Perotto et al. 2018; Wylie et al. 2017).

Cucurbit yellow stunting disorder virus Taxonomic position Genus: Crinivirus

(CYSDV)

Family: Closteroviridae

CYSDV infection in plants of Cucurbita maxima was reported from Sudan (Mohammed et al. 2014). The virus-infected pumpkin plants exhibit symptoms of interveinal chlorosis, yellowing, and brittleness of lower leaves. The virus is transmitted by a whitefly vector, Bemisia tabaci, in a semi-persistent manner. The virus is not mechanically sap-transmissible. For more details of CYSDV, refer to Cucumis melo.

Lettuce infectious yellows virus Taxonomic position Genus: Crinivirus

(LIYV)

Family: Closteroviridae

LIYV infection in plants of Cucurbita maxima was reported from Texas (USA) (Halliwell and Johnson 1992). The virus-infected pumpkin plants exhibit various degrees of mosaic symptoms, and fruits do not ripen completely and plants are stunted. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a semi-persistent manner. The virus is not mechanically sap-transmissible. For more details of LIYV, refer to Lactuca sativa.

Papaya ringspot virus Taxonomic position Genus: Potyvirus

(PRSV)

Family: Potyviridae

PRSV pathotype W was previously known as Watermelon mosaic virus-1. PRSV-W is known to infect plants of Cucurbita maxima in India, Tunisia, Egypt, Nigeria, and Sudan (Thomas 1980; Biswas and Varma 2006; Krishna Kumar et al. 2007; Omar et al. 2011; Mohammed et al. 2012; Nagendran et al. 2017). The virus induces greenish mosaic, blistering, and malformation of leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PRSV, refer to Carica papaya.

C

734

Pea streak virus Taxonomic position Genus: Carlavirus

Cucurbita maxima (Pumpkin)

(PeSV)

Family: Betaflexiviridae

PeSV infection in plants of Cucurbita maxima was reported from Saudi Arabia (Al-Shahwan et al. 2017). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of PeSV, refer to Pisum sativum.

Pumpkin yellow mosaic virus

(PuYMV)

Synonyms Pumpkin yellow mosaic Malaysia virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution PuYMV infection in plants of Cucurbita maxima was reported from Malaysia and India (Singh et al. 1991; Tsai et al. unpublished - EF197941). Symptoms and host(s) The virus-infected pumpkin plants exhibit yellow mosaic symptoms. Transmission The transmission of the PuYMV has not been investigated. It is likely that, in common with other begomoviruses, PuYMV is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of PuYMV has not been investigated. In common with all geminiviruses the virions of PuYMV will likely be geminate (twinned quasi-icosahedra). With only a single isolate of PuYMV having been characterised, the precise nature of the genome (whether monopartite, monopartite betasatellite-associated or biapartite) is unclear. The characterised genome/genomic component of PuYMV is 2724 nt (EF197941 = NC_010946) (Briddon 2001; Tsai et al. 2006; Brown et al. 2015; Zerbini et al. 2017). The characterised genome/genomic component of PuYMV encodes the six genes typically encoded by monopartite begomoviruses and the DNA A component of bipartite begomoviruses. The expression and function of the genes have not been investigated.

Red clover vein mosaic virus Taxonomic position Genus: Carlavirus

(RCVMV)

Family: Betaflexiviridae

Cucurbita maxima (Pumpkin)

735

RCVMV infection in plants of Cucurbita maxima was reported from Saudi Arabia (Al-Shahwan et al. 2017). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation to a number of hosts. For more details of RCVMV, refer to Trifolium spp.

Squash leaf curl China virus

(SLCCNV)

Synonyms Pumpkin yellow vein mosaic virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Pumpkin yellow vein mosaic virus (PYVMV) was an isolate of Squash leaf curl China virus (SLCCNV) (Maruthi et al. 2007). SLCCNV infection in plants of Cucurbita maxima was reported from Thailand, Pakistan, and India (Maruthi et al. 2003; Muniyappa et al. 2003; Ito et al. 2008; Singh et al. 2008, 2009; Tiwari et al. 2011; Hamsa et al. 2016). The virus-infected pumpkin plants exhibit yellow chlorotic spot, mosaic, and leaf curling symptoms. Flower and fruits of the severely infected plants were distorted. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not mechanically sap-transmissible. For more details of SLCCNV, refer to Cucurbita pepo.

Squash mosaic virus

(SqMV)

Synonyms Pumpkin mosaic virus Taxonomic position Genus: Comovirus

Family: Secoviridae

SqMV infection in plants of Cucurbita maxima was reported from the Czech Republic, Western Samoa, Trinidad, and Turkey (Sevik and Toksoz 2008; Pearson and Liayanage 1997; Svoboda and LeisovaSvobodova 2011; Chinnaraja et al. 2016). The virus-infected pumpkin plants exhibit yellow mosaic and leaf deformation symptoms. The virus is transmitted by beetle vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SqMV, refer to Cucurbita pepo.

Tobacco mild green mosaic virus Taxonomic position Genus: Tobamovirus

(TMGMV)

Family: Virgaviridae

TMGMV infection in plants of pumpkin (Cucurbita moschata) was reported from China (Tang et al. 2017). The virus-infected pumpkin plants exhibit mosaic and leaf distortion symptoms. The virus is transmissible by mechanical sap-inoculation, and has a wide host range. The virus is transmissible by grafting and also by contact between plants. For more details of TMGMV, refer to Nicotiana tabacum.

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736

Cucurbita maxima (Pumpkin)

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Cucurbita maxima was reported from Oklahoma (Abdalla et al. 2012). The virus-infected pumpkin plants exhibit symptoms of chlorotic spots, systemic ringspot, severe leaf deformation, mottling, and stunting. The virus is transmitted through nematode vectors in a nonpersistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato chlorosis virus Taxonomic position Genus: Crinivirus

(ToCV)

Family: Closteroviridae

ToCV infection in plants of pumpkin (Cucurbita moschata) was reported from China (Sun et al. 2016a). The virus-infected plants exhibit chlorotic spot symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a semi-persistent manner. For more details of ToCV, refer to Solanum lycopersicum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV was detected in plants of Cucurbita maxima in Andhra Pradesh, India (Sarovar et al. 2010). The virus-infected pumpkin plants exhibit chlorosis and necrotic symptoms on the leaves, stem and inflorescence, stunting, and wilting of plants. The virus is transmitted by the thrips vectors. The virus present in/on pollen, entering in to the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Cucurbita maxima or Cucurbita moschata was reported from India and Italy (Phaneendra et al. 2012; Parrella et al. 2018). The virus-infected pumpkin plants exhibit symptoms of marginal rolling and curling of leaves and yellow patches on lamina. The affected plants were stunted and produced less flowers and fruits. The virus is transmitted by a whitefly vector, Bemisia tabaci, in a circulative non-propagative manner, and also by grafting. For more details of ToLCNDV, refer to Solanum lycopersicum.

Cucurbita maxima (Pumpkin)

737

Tomato leaf curl Palampur virus Taxonomic position Genus: Begomovirus

(ToLCPalV)

Family: Geminiviridae

In northern India, ToLCPalV was identified to be associated with yellowing and curling disease of pumpkin (Cucurbita moschata) (Namrata et al. 2010). The virus is transmitted by a whitefly vector, Bemisia tabaci, in a circulative non-propagative manner, and also by grafting. For more details of ToLCPalV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of pumpkin (Cucurbita moschata) was reported from China (Sun et al. 2016b). The virus-infected pumpkin plants exhibit leaf mottling, crinkling, and mosaic symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sapinoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Cucurbita maxima was reported from Japan (Shahid et al. 2015). The virus-infected pumpkin plants exhibit foliar yellowing and curling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV infection in plants of Cucurbita maxima occurs wherever the crop is grown (Sako et al. 1975; Singh et al. 1991; Fletcher et al. 2000; Wakman et al. 2002). The virus-infected pumpkin plants exhibit systemic mosaic of green and yellow green color, vein-banding, and sometimes malformation. The infected plants produced abnormal flowers with underdeveloped or incompletely opened petals. The aphid vector, Myzus persicae, transmits the virus in a non-persistent manner, and also through mechanical sap-inoculation. The virus is not transmitted through pumpkin seeds. For more details of WMV, refer to Citrullus lanatus.

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Zucchini tigre mosaic virus Taxonomic position Genus: Potyvirus

(ZTMV)

Family: Potyviridae

ZTMV infection in plants of Cucurbita maxima was reported from China (Xiao et al. 2016). The virusinfected pumpkin plants exhibit blistering, mosaic mottling, chlorotic, and dark-green vein-banding symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ZTMV, refer to Cucurbita pepo.

Zucchini yellow mosaic virus Taxonomic position Genus: Potyvirus

(ZYMV)

Family: Potyviridae

ZYMV infection in plants of Cucurbita maxima was worldwide in distribution (Providentii et al. 1984; Ohtsu and Sako 1985; Fletcher 1996; Pearson and Liayanage 1997; Fletcher et al. 2000; Gracia 2000; Auger et al. 2004; Muller et al. 2006; Vucurovic et al. 2012; Romay et al. 2014; Chinnaraja et al. 2016; Nagendran et al. 2016; Karthikeyan et al. 2017). The virus-infected pumpkin leaves show mosaic and mottling symptoms which were received for disease diagnosis. Leaf symptoms in the field included yellow vein-banding, chlorotic yellow mosaic with distortion and blister of leaves, and sometimes a “fan leaf” effect. Some plants were stunted. Fruit symptoms may include pimpling, blister, and reduced size. The virus is transmitted by a wide range of aphid species of which the green peach (Myzus persicae) and melon (Aphis gossypii) aphids are the most important. The virus is mechanically saptransmissible. The virus is not seed-borne in pumpkin seeds (Gleason and Provvidenti 1990). For more details of ZYMV, refer to Cucurbita pepo.

References Abdalla OA, Bruton BD, Fish WW, Ali A (2012) First confirmed report of Tobacco ringspot virus in cucurbit crops in Oklahoma. Plant Dis 96:1705 Ali A, Hussain A, Ahmad M (2014) Occurrence and molecular characterization of Cucumber green mottle mosaic virus in cucurbit crops of KPK, Pakistan. Braz J Microbiol 45:1247–1253 Al-Shahwan IM, Abdalla OA, Al-Saleh MA, Amer MA (2017) Detection of new viruses in alfalfa, weeds and cultivated plants growing adjacent to alfalfa fields in Saudi Arabia. Saudi J Biol Sci 24(6):1336–1343 Auger J, Martinez C, Esterio M, Prieto H (2004) Identification and transmission of strains of 24 zucchini yellow mosaic virus (ZYMV) causal agents of fruit deformation in squash (Cucurbita maxima) in Chile. Fitopatologia 39:133–143 Biswas C, Varma A (2006) Characterization of virus from pumpkin as an isolate of PRSV-W. Indian Phytopathol 59:101–104 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Idris AM, Olsen MW, Miller ME, Isakeit T, Anciso J (2000) Cucurbit leaf curl virus, a new whitefly transmitted geminivirus in Arizona, Texas and Mexico. Aust Plant Dis Notes 84:809 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619

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Chinnaraja C, Ramkissoon A, Rajendran R, Tony ST, Ramsubhag A, Jayaraj J (2016) First report of Zucchini yellow mosaic virus and Squash mosaic virus infecting cucurbits in Trinidad. Plant Dis 100:866 Fletcher JD (1996) Zucchini yellow mosaic virus in buttercup squash – a new record in New Zealand. Aust Plant Pathol 25:142 Fletcher JD, Wallace AR, Rogers BT (2000) Potyviruses in New Zealand buttercup squash (Cucurbita maxima Duch.): yield and quality effects of ZYMV and WMV 2 virus infections. N Z J Crop Hortic Sci 28:17–26 Gleason ML, Provvidenti R (1990) Absence of transmission of Zucchini yellow mosaic virus from seeds of pumpkin. Plant Dis 74:828 Gracia O (2000) First report of Zucchini yellow mosaic virus in Argentina. Aust Plant Dis Notes 84:371 Halliwell RS, Johnson JD (1992) Lettuce infectious yellows virus infecting watermelon, cantaloupe, honey dew melon, squash and cushaw in Texas. Plant Dis 76:643 Hamsa S, Girija D, Nazeem PA, Mathew SK, Deepa K, Balakrishnan S, Sumbula V (2016) Molecular characterization of geminivirus causing yellow vein mosaic in pumpkin. Int J Sci Environ Technol 5:133–147 Hattab MM, Guathier N, Zouba A (2009) Biological and molecular characterization of the Cucurbit aphid-borne yellows virus affecting cucurbits in Tunisia. Plant Dis 93:1065–1072 Huang LH, Tseng HH, Li JT, Chen TC (2010) First report of Cucurbit chlorotic yellows virus infecting Cucurbits in Taiwan. Plant Dis 94:1168 Ito T, Ogawa T, Samretwanich K, Sharma P, Ikegami M (2008) Yellow leaf curl disease of pumpkin in Thailand is associated with Squash leaf curl China virus. Plant Pathol 57:766 Karthikeyan G, Mano Ranjitham SK, Nagendran K (2017) Zucchini yellow mosaic virus – an emerging threat to Cucurbitaceous vegetables cultivation. In: 26th annual conference of Indian virological society (VIROCON 3017), Mangaluru, India, p 106 Knierim D, Deng TC, Tsai WS, Green SK, Kenyon L (2010) Molecular identification of three distinct Polerovirus species and a recombinant Cucurbit aphid-borne yellows virus strain infecting cucurbit crops in Taiwan. Plant Pathol 59:991–1002 Krishna Kumar NK, Kalleshwaraswamy CM, Saroja S, Krishna Reddy M, Venugopalan R (2007) Transmission and epidemiology of Papaya ringspot virus – W (PRSV-W) infecting pumpkin. In: 10th international plant virus epidemiology symposium 15–19 October 2007, ICRISAT, India, p 41. (Abstract) Kumar A, Jailani AAK, Roy A, Mandal B (2017) The occurrence, biology and genomic properties of tobamoviruses infecting crop plants in India. In: Mandal B, Rao GP, Baranwal VK, Jain RK (eds) A century of plant virology in India. Springer Nature, Singapore, pp 429–443. ISBN 978-981-10-5671-0 Lecoq H, Bourdin D, Wipf-Scheibel C, Bon M, Lot H, Lemaire O, Herrbach E (1992) A new yellowing disease of cucurbits caused by a Luteovirus, Cucurbit aphid-borne yellows virus. Plant Pathol 41:749–761 Maruthi MN, Colvin J, Briddon RW, Bull SE, Muniyappa V (2003) Pumpkin yellow vein mosaic virus: a novel Begomovirus infecting cucurbits. J Plant Pathol 85:64 Maruthi MN, Rekha AR, Muniyappa V (2007) Pumpkin yellow vein mosaic disease is caused by two distinct begomoviruses: Complete viral sequences and comparative transmission by an indigenous Bemisia tabaci and the introduced B-biotype. Bull OEPP/EPPO Bull 37(2):412–419 Mohammed H, Manglli A, Zicca S, El Hussein A, Mohamed M, Tomassoli L (2012) First report of Papaya ringspot virus in pumpkin in Sudan. New Dis Rep 26:26 Mohammed HS, Zicca S, Manglli A, Mohamed ME, El Siddig MA, Tomassoli L, El Hussein AA (2014) Identification and phylogenetic analysis of common pumpkin viruses in Sudan. J Plant Pathol 96:77–84 Mukhopadhayay S, Saha K (1968) Transmission of Cucumis virus (Cucumber mosaic virus) through seeds of Cucurbita maxima L. Sci Cult 34:436–437 Muller C, Bröther H, von Bargen S, Büttner C (2006) Zucchini yellow mosaic virus incidence and sources of virus infection in field-grown cucumbers and pumpkins in the Spreewald, Germany. J Plant Dis Protect 113:252–258 Muniyappa V, Maruthi MN, Babitha CR, Colvin J, Briddon RW, Rangaswamy KT (2003) Characterisation of pumpkin yellow vein mosaic virus from India. Ann Appl Biol 142:323–331 Nagendran K, Priyanka R, Keerthana U, Mohankumar S, Karthikeyan G (2016) First report of Zucchini yellow mosaic virus on Cucurbita moschata in India. J Plant Pathol 98:173 Nagendran K, Mohankumar S, Aravintharaj R, Balaji CG, Manoranjitham SK, Singh AK, Rai AB, Singh B, Karthikeyan G (2017) The occurrence and distribution of major viruses infecting cucurbits in Tamil Nadu state, India. Crop Prot 99:10–16 Namrata J, Saritha RK, Datta D, Singh M, Dubey RS, Rai AB, Rai M (2010) Molecular characterization of Tomato leaf curl Palampur virus and Pepper leaf curl betasatellite Naturally Infecting Pumpkin (Cucurbita moschata) in India. Indian J Virol 21:128–132 Ohtsu Y, Sako N (1985) Zucchini yellow mosaic virus isolated from pumpkin in Milyako and Yaeyama islands, Okinawa, Japan. Ann Phytopathol Soc Jpn 51:234–237

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Omar AF, El-Kewey SA, Sidaros SA, Shimaa AK (2011) Egyptian isolates of Papaya ringspot virus form a molecularly distinct clade. J Plant Pathol 93:1224 Parrella G, Troiano E, Formisano G, Accotto GP, Giorgini M (2018) First report of Tomato leaf curl New Delhi virus associated with severe mosaic of pumpkin in Italy. Plant Dis 102:459 Pearson MN, Liayanage AS (1997) Records of cucurbit viruses infecting vegetable crops in Western Samoa. Australas Plant Pathol 28:188–191 Perotto MC, Pozzi EA, Celli MG, Luciani CE, Mitidieri MS, Conci VC (2018) Identification and characterization of a new Potyvirus infecting cucurbits. Arch Virol 163(3):719–724 Phaneendra C, Rao KRSS, Jain RK, Mandal B (2012) Tomato leaf curl New Delhi virus is associated with pumpkin leaf curl: a new disease in northern India. Indian J Virol 23(1):42–45 Providentii R, Gonsalves D, Humaydan HS (1984) Occurrence of Zucchini yellow mosaic virus in cucurbits from Connecticut New York, Florida, and California. Plant Dis 68:443–446 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Romay G, Lecoq H, Desbiez C (2014) Cucurbit crops and their viral diseases in Latin America and the Caribbean Islands: a review. J Plant Pathol 96(2):227–242 Sako N, Nomura M, Nonaka F (1975) Two isolates of watermelon mosaic virus isolated from pumpkin leaves. Bull Fac Agric Saga Univ 39:1–10 Sarovar B, Prasad YS, Sai Gopal DVR (2010) Detection of Tobacco streak virus by immunocapture-reverse transcriptasepolymerase chain reaction and molecular variability analysis of a part of RNA3 of sunflower, gherkin, and pumpkin from Andhra Pradesh, India. Sci Asia 36:194–198 Sevik MA, Toksoz Y (2008) Occurrence of Squash mosaic virus (SqMV) infecting pumpkin and squash in Samsun, Turkey. J Turk Phytopathol 37:15–25 Shahid MS, Ikegami M, Briddon RW, Natsuaki KT (2015) Characterization of Tomato yellow leaf curl virus and associated alphasatellite infecting Cucurbita maxima in Japan. J Gen Plant Pathol 81:92–95 Singh BR, Gupta SP, Roy PS (1991) Prevalence of viral diseases on pumpkin in Uttar Pradesh. Indian J Virol 7:190–191 Singh R, Raj SK, Prasad V (2008) Molecular characterization of a strain of Squash leaf curl China Virus from North India. J Phytopathol 156:222–228 Singh AK, Mishra KK, Chattopadhyay B, Chakraborty S (2009) Biological and molecular characterization of a begomovirus associated with yellow mosaic vein mosaic disease of pumpkin from Northern India. Virus Genes 39:359–370 Sun XH, Gao LL, Wang SL, Wang CL, Yang YY, Wang XY, Zhu XP (2016a) First report of Tomato spotted wilt virus infecting pumpkin in China. J Plant Pathol 98:687 Sun XH, Gao LL, Wang CL, Wang SL, Zhao J, Qiao N, Zhu XP (2016b) Natural occurrence of Tomato chlorosis virus in pumpkin in China. Plant Dis 100:2543 Svoboda J, Leisova-Svobodova L (2011) First report of Squash mosaic virus in ornamental pumpkin in the Czech Republic. Plant Dis 95:1321 Tang Q-J, Liu M-Y, Li X, Zhao S-Q, Dai L-Y (2017) Pumpkin: a new natural host of Tobacco mild green mosaic virus in China. Plant Dis 101:1063 Thomas W (1980) Watermelon mosaic virus, the cause of a serious disease of cucurbits in Cook Islands. N Z J Exp Agric 8:309–312 Tiwari AK, Snehi SK, Singh R, Raj SK, Rao GP, Sharma PK (2011) Molecular identification and genetic diversity among six Begomovirus isolates affecting cultivation of cucurbitaceous crops in Uttar Pradesh, India. Arch Phytopathol Plant Protect 45:62–72 Vucurovic A, Bulajic A, Stankovic I, Ristic D, Berenji J, Jovic J, Krstic B (2012) Non-persistently aphid-borne viruses infecting pumpkin and squash in Serbia and partial characterization of Zucchini yellow mosaic virus isolates. Eur J Plant Pathol 133:935–947 Wakman W, Kontong MS, Teakle DS, Persley DM (2002) Watermelon mosaic virus of pumpkin (Cucurbita maxima) from Sulawesi: identification, transmission and host range. Indones J Agric Sci 3:33–36 Wintermantel WM (2004) Pumpkin (Cucurbita maxima and C. pepo), a new host of Beet pseudo yellows virus in California. Plant Dis 88:82 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Xiao L, Li YY, Tan GL, Lan PX, Zhong L, Liu Y, Li R, Li F (2016) First report of Zucchini tigre mosaic virus infecting several cucurbit plants in China. Plant Dis 100:1253 Yardimci N, Ozgonen H (2007) First report of Cucurbit aphid-borne yellows virus in Turkey. Aust Plant Dis Notes 2:59 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zitikaite I, Staniulis J, Urbanavicien_e L, Zizyt M (2011) Cucumber mosaic virus identification in pumpkin plants. Zemdirbyste Agric 98:421–426

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Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow) Family: Cucurbitaceae

Vegetable

Algerian watermelon mosaic virus Taxonomic position Genus: Potyvirus

(AWMV)

Family: Potyviridae

Geographical distribution AWMV infection in plants of Cucurbita pepo was reported from Tunisia and Algeria (Yakoubi et al. 2008). Symptoms and host(s) The virus-infected zucchini plants exhibit mosaic symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically saptransmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome is a single molecule of linear, positive-sense single-stranded RNA of 9842 nt (EU410442 = NC_010736) (Yakoubi et al. 2008; Revers and Garcia 2015; Wylie et al. 2017).

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Cucurbita pepo was reported from the USA (Provvidenti and Uyemoto 1973). The virus-infected zucchini plants exhibit chlorotic leaf spotting symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Beet pseudoyellows virus Taxonomic position Genus: Crinivirus

(BPYV)

Family: Closteroviridae

BPYV infection in plants of Cucurbita pepo was reported from California and Costa Rica (Wintermantel 2004; Hammond et al. 2005). The virus-infected zucchini plants exhibit yellowing and leaf chlorosis symptoms. The virus is transmitted by the whitefly vector, Trialeurodes

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vaporariorum, in a semi-persistent manner. The virus is not transmissible by mechanical sap-inoculation, and also not by contact between plants. For more details of BPYV, refer to Beta vulgaris.

Capsicum chlorosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(CaCV)

Family: Tospoviridae

CaCV infection in plants of Cucurbita pepo was reported from China (Sun et al. 2018). The virusinfected Summer squash plants exhibit fruits had chlorosis and rugged symptoms. The virus is transmitted by thrips vectors in a persistent, propagative manner, and also by mechanical sap inoculation. For more details of CaCV, refer to Capsicum annuum.

Chickpea chlorotic dwarf virus Taxonomic position Genus: Mastrevirus

(CpCDV)

Family: Geminiviridae

CpCDV infection in plants of Cucurbita pepo was reported from Egypt (Fahmy et al. 2015). The virusinfected zucchini plants exhibit severe chlorosis and stunting symptoms. The virus is transmitted by leafhopper vectors in a persistent, circulative, and non-propagative manner. The virus is not transmissible by mechanical inoculation. For more details of CpCDV, refer to Cicer arietinum.

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

CGMMV infection in plants of Cucurbita pepo was reported from Pakistan and China (Lovisolo 1981; Zhang et al. 2009; Ali et al. 2014). The virus-infected zucchini plants exhibit symptoms of light yellow green spots and vein-clearing on young leaves to chlorotic mottling, plant stunting, and necrosis at later stages of growth. The virus is transmissible by mechanical sap-inoculation, and also by contact between plants. Transmitted through contaminated irrigation water or nutrient solutions has also been reported. For more details of CGMMV, refer to Cucumis sativus.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV occurs wherever Cucurbita pepo plants are grown (Sammons et al. 1989; Wong et al. 1999; Bostan et al. 2002; Dukic et al. 2002; Krstic et al. 2002; Bashir et al. 2006; Ozaslan et al. 2006; Jossey and Babadoost 2008; Kim et al. 2010; Vucurovic et al. 2011; Sharma et al. 2013; Tiwari and Gao 2014; Hasiow-Jaroszewska et al. 2017). The virus-infected zucchini plants exhibit symptoms such as severe epinasty and downward bending of the petiole and leaf surface along with leaf reduction, which are

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common in early season infection of squash. The virus is transmitted by several aphid species in a nonpersistent manner, and also by mechanical sap-inoculation to a large number of host plants. The virus is seed-transmitted in Cucurbita pepo seeds which was up to 5.1% (Sharma and Chohan 1973; Tobias et al. 2008; Sevik and Balkaya 2015). For more details of CMV, refer to Cucumis sativus.

Cucumber vein yellowing virus Taxonomic position Genus: Ipomovirus

(CVYV)

Family: Potyviridae

CVYV-infected plants of Cucurbita pepo were reported from the Mediterranean Basin, Spain, southern Portugal, and Tunisia (Gil-Salas et al. 2011; Louro et al. 2004; Yakoubi et al. 2007). The virus-infected zucchini plants show vein-clearing and mottling on leaves. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CVYV, refer to Cucumis sativus.

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(CABYV)

Family: Luteoviridae

CABYV infection in plants of Cucurbita pepo was reported from China, Algeria, Greece, Turkey, Lebanon, Libya, Spain, Italy, Taiwan, Tunisia, Egypt, Montenegro, Saudi Arabia, and the Czech Republic (Lecoq et al. 1992, 1994; Abou-Jawdah et al. 1997; Hattab et al. 2005, 2009; Kassem et al. 2007; Yardimci and Ozgonen 2007; Xiang et al. 2008; Knierim et al. 2010; Svoboda et al. 2011, 2015; Omar and Bagdady 2012; Al-Saleh et al. 2015; Zindovic et al. 2017). The virus-infected zucchini plants exhibit yellowing of the older leaves, and leaves are thickened and brittle. A wide range of symptom intensity exists depending on the cultivars, varying from a yellowing limited to a few older leaves to a complete discoloration of plants. The virus infection does not affect fruit quality but rather induces flower abortions and reduces the number of fruit per plant. CABYV has a wide host range infecting many cucurbits (Lecoq et al. 1992). The virus is transmitted by aphid vectors, Myzus persicae, M. euphorbiae, and Aphis gossypii, in a circulative, non-propagative manner (Lecoq et al. 1992). The virus is not transmissible by mechanical sap-inoculation. For more details of CABYV, refer to Cucumis melo.

Cucurbit chlorotic yellows virus

(CCYV)

Taxonomic position CCYV is a tentative member of the genus Crinivirus and family Closteroviridae CCYV infection in plants of Cucurbita pepo was reported from Greece, Iran, and Taiwan (Huang et al. 2010; Bananej et al. 2013). The virus-infected zucchini plants exhibit symptoms of chlorosis, interveinal chlorotic spots on lower leaves, and bright yellow color or severe yellowing on older leaves. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a semi-persistent manner. For more details of CCYV, refer to Cucumis melo.

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Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow)

Cucurbit leaf crumple virus

(CuLCrV)

Synonyms Cucurbit leaf curl virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

CuLCrV infection in plants of Cucurbita pepo was reported from Florida, Arizona, Mexico, and California (Gilbertson 2002; Akad et al. 2008). The virus-infected zucchini plants exhibit interveinal chlorosis and crumpled, curled, thickened, and rounded leaves. Infected fruit is streaked with green, making them unmarketable. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not mechanically sap-transmissible. For more details of CuLCrV, refer to Cucumis melo.

Cucurbit mild mosaic virus Taxonomic position Genus: Fabavirus

(CuMMV)

Family: Secoviridae

Geographical distribution CuMMV infection in plants of Cucurbita pepo was reported from China (Dong et al. 2012). Symptoms and host(s) The virus-infected zucchini plants exhibit inconspicuous mosaic symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically saptransmissible. Virion properties and genome The virions are isometric, non-enveloped of two types but similar in size 25–30 nm in diameter, and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is a bipartite, linear positive-sense, single-stranded RNA. RNA1 consists of 5870 nt (EU881936) and RNA2 is 3294 nt (EU881937) (Ikegami and Sharma 2011; Sanfacon et al. 2009; Dong et al. 2012; Sanfacon 2015; Thompson et al. 2017).

Cucurbit yellow stunting disorder virus Taxonomic position Genus: Crinivirus

(CYSDV)

Family: Closteroviridae

CYSDV infection in plants of Cucurbita pepo was reported from UAE, Florida, Mexico, Saudi Arabia, and Tunisia (Hassan and Duffus 1991; Abou-Jawdah et al. 2000; Brown et al. 2007; Yakoubi et al. 2007; Polston et al. 2008; Al-Saleh et al. 2015). The virus-infected zucchini plants show severe yellowing symptoms that started as interveinal mottle on the older leaves and developed into severe

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yellowing symptoms as the leaves became older. The virus is transmitted by both the B and Q biotypes of B. tabaci in a semi-persistent manner (Berdiales et al. 1999; McGrath 2004). The virus is not mechanically sap-transmissible. For more details of CYSDV, refer to Cucumis melo.

Lettuce infectious yellows virus Taxonomic position Genus: Crinivirus

(LIYV)

C

Family: Closteroviridae

LIYV infection in plants of Cucurbita pepo was reported from the USA (Cohen et al. 1992; Halliwell and Johnson 1992). The virus-infected zucchini plants exhibit silverleaf disorder symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a semi-persistent manner (Cohen et al. 1992). The virus is not mechanically sap-transmissible and is not seed-transmitted. For more details of LIYV, refer to Lactuca sativa.

Melon severe mosaic orthotospovirus Taxonomic position Genus: Orthotospovirus

(MSMV)

Family: Tospoviridae

MSMV infection in plants of Cucurbita pepo was reported from Mexico (Ciuffo et al. 2009). The virus is transmitted by a thrips vector Frankliniella occidentalis in a persistent propagative manner, and also by mechanical sap-inoculation to a number of herbaceous hosts. For more details of MSMV, refer to Cucumis melo.

Moroccan watermelon mosaic virus Taxonomic position Genus: Potyvirus

(MWMV)

Family: Potyviridae

MWMV infection in plants of zucchini squash (Cucurbita pepo) was reported from France, Greece, and Italy (Roggero et al. 1998; Lecoq et al. 2008; Malandraki et al. 2014). The virus-infected zucchini plants show severe mosaic and blisters and deformation of leaves, as well as fruit deformations. The virus is transmitted by the aphid vector (Myzus persicae) in a non-persistent manner, and also by mechanical sap-inoculation only to species belonging to the Cucurbitaceae family. For more details of MWMV, refer to Citrullus lanatus.

Ourmia melon virus Taxonomic position Genus: Ourmiavirus

(OuMV)

Family: Botourmiaviridae

OuMV infection in plants of Cucurbita pepo was reported from Northern Iran (Gholamalizadeh et al. 2008). The virus-infected zucchini plants exhibit mosaic, yellowing, chlorotic spot, and irregular

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Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow)

ringspot symptoms. No vector has been identified. The virus is mechanically sap-transmissible to a number of dicotyledon plants. For more details of OuMV, refer to Cucumis melo.

Papaya mosaic virus Taxonomic position Genus: Potexvirus

(PapMV)

Family: Alphaflexiviridae

PapMV infection in plants of Cucurbita pepo was reported from Mexico (Noa-Carrazana and SilvaRosales 2001). The virus is transmitted by means not involving a vector. The virus is mechanically saptransmissible. For more details of PapMV, refer to Carica papaya.

Papaya ringspot virus Taxonomic position Genus: Potyvirus

(PRSV)

Family: Potyviridae

PRSV (pathotype W) infection in plants of Cucurbita pepo was reported from the USA, Cuba, Poland, China, India, and South Africa (Sammons et al. 1989; Jossey and Babadoost 2008; HasiowJaroszewska et al. 2010; Tiwari and Gao 2014; Ibaba et al. 2015; Rodriguez-Martinez et al. 2015; Mahesha and Krishna Reddy 2016; Cheng et al. 2017). The virus-infected zucchini plants show leaf mosaic and marbled fruit symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PRSV, refer to Carica papaya.

Pepo aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(PABYV)

Family: Luteoviridae

Geographical distribution PABYV infection in plants of Cucurbita pepo was reported from South Africa, Cote d’Ivoire, Mali and Thailand (Knierim et al. 2014; Kone et al. 2015; Ibaba et al. 2017). Transmission The virus is transmitted by a number of aphid species in a circulative and non-propagative manner. The virus is not sap or seed transmitted. Virion properties and genome The virions are 25–30 nm in diameter, hexagonal in outline and have no envelope. The genome contains a single molecule of positive-sense ssRNA as its genome of 5813 nt (KU315178 = NC_030225; KU315177). The 30 terminus has neither a poly (A) tract nor a tRNA-like structure. The 50 terminus has a genome-linked protein (of about 17 kDa). The genome contains six ORFs (Knierim et al. 2014; Ibaba et al. 2017).

Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow)

Pepper golden mosaic virus Taxonomic position Genus: Begomovirus

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(PepGMV)

Family: Geminiviridae

PepGMV infection in plants of Cucurbita pepo var. moschata (squash) and Cucurbita pepo was reported from Costa Rica (Castro et al. 2013). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of PepGMV, refer to Capsicum annuum.

Squash chlorotic leaf spot virus Taxonomic position Genus: Torradovirus

(SCLSV)

Family: Secoviridae

Geographical distribution SCLSV infection in plants of Cucurbita pepo var. moschata (squash) was reported from Sudan (Lecoq et al. 2016). Symptoms and host(s) The virus-infected summer squash plants exhibit chlorotic leaf spot symptoms. Transmission The virus is transmitted by the greenhouse whitefly (Trialeurodes vaporariorum) and silverleaf whitefly (Bemisia tabaci) in a semi-persistent manner (Verbeek et al. 2014). The virus is also mechanically transmissible to squash (Cucurbita moschata) and other cucurbits. Virion properties and genome The virions are isometric, non-enveloped of two types but similar in size 25–30 nm in diameter, and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is a bipartite, linear positive-sense, single-stranded RNA. RNA1 consists of 7045 nt (KU052530) and RNA2 of 4730 nt (KU052531) (Sanfacon et al. 2009; Sanfacon 2015; van der Vlugt et al. 2015; Lecoq et al. 2016; Thompson et al. 2017).

Squash leaf curl China virus

(SLCCNV)

Synonyms Pumpkin yellow vein mosaic virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution SLCCNV infection in plants of Cucurbita pepo was reported from China, India, Vietnam, and the Philippines (Hong et al. 1995; Kon et al. 2003; Tahir et al. 2010; Saritha et al. 2011).

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Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow)

Symptoms and host(s) The virus-infected summer squash plants exhibit yellow spots or patches on leaves, which coalesce turning the leaf completely yellow with a few green islands. Curling or deformation of leaves is associated with the yellowing and mottling symptoms. The virus has been isolated from a range of cucurbit species including pumpkin/squash/zucchini (Cucurbita pepo, C. maxima) and wax gourd (Benincasa hispida). Infected plants show a yellow vein mosaic or yellow patches (mottling) and leaf curling. In some cases the leaves may be almost entirely chlorotic with only a few dark green patches. Transmission The virus has been transmitted, using the vector of begomoviruses the whitefly Bemisia tabaci, from field-infected C. pepo to healthy C. pepo plants (Singh et al. 2008). However, the mechanism of transmission of SLCCNV has not been investigated. It is likely that, in common with other begomoviruses, SLCCNV is transmitted by the whitefly in a circulative, non-propagative manner. Virion properties and genome The virion structure of SLCCNV has not been investigated. In common with all geminiviruses, the virions of SLCCNV are likely geminate (twinned quasi-icosahedra). The genome of SLCCNV is a bipartite consisting of two circular, single-stranded DNA components. DNA-A contains 2736 nt (AF509743 = NC_007339) and DNA-B of 2719 nt (AF509742 = NC_007338) (Briddon 2001; Kon et al. 2003; Revill et al. 2003; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the Old World, the DNA-A component of SLCCNV encodes six genes, two in the virion-sense and four in the complementarysense, and the DNA-B encodes two genes, one in each orientation. The expression and function of these genes have not been investigated for SLCCNV.

Squash leaf curl Philippines virus Taxonomic position Genus: Begomovirus

(SLCuPV)

Family: Geminiviridae

Geographical distribution SLCuPV infection in plants of Cucurbita pepo was reported from the Philippines and Taiwan (Kon et al. 2003; Tsai et al. 2007). Symptoms and host(s) The virus-infected summer squash plants exhibit leaf curling symptoms. The virus has been isolated from squash (Cucurbita moschata), pumpkin (Cucurbita pepo), winter melon (Benincasa hispida), and chayote (Sechium edule). Infected cucurbit plants exhibit leaf curling, blistering, and yellowing symptoms. Transmission The transmission of the SLCuPV has not been investigated. It is likely that, in common with other begomoviruses, SLCuPV is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible.

Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow)

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Virion properties and genome The virion structure of SLCuPV has not been investigated. In common with all geminiviruses, the virions of SLCuPV are likely geminate (twinned quasi-icosahedra). The genome of SLCuPV is a bipartite consisting of two circular, single-stranded DNA components. DNA-A contains 2739 nt (AB085793 = NC_005845; EU479710) and DNA-B 2705 nt (AB085794; EU479711) (Briddon 2001; Kon et al. 2003; Tsai et al. 2007; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the Old World, the DNA-A component of SLCuPV encodes six genes, two in the virion-sense and four in the complementary-sense, and the DNA-B encodes two genes, one in each orientation. The expression and function of these genes have not been investigated for SLCuPV.

Squash leaf curl virus Taxonomic position Genus: Begomovirus

(SLCuV)

Family: Geminiviridae

Geographical distribution SLCuV was originally identified on squash (Cucurbita pepo) showing leaf curl disease in California, USA (Flock and Mayhew 1981). The virus spreads in Asia, Africa, North America, Central America, Egypt, Jordan, Palestine, Israel and the Caribbean (Dodds et al. 1984; Idris et al. 2006; Al-Musa et al. 2008; Abudy et al. 2010; Ali-Shtayeh et al. 2010, 2014; Sufrin-Ringwald and Lapidot 2011; Sobh et al. 2012). Symptoms and host(s) Characteristic symptoms of SLCuV on squash include severe chlorotic foliar mosaic and mottling, leaf curling, twisting, malformation, yellowing, leaves with shorter petioles, sterility, stunting of plats, and, in some cultivars, distortion of fruit. The virus causes severe systemic stunting and leaf curling in most cucurbits. In pumpkin, squash, and watermelon, a severe, chlorotic foliar mosaic or mottle accompanies leaf curling and stunting symptoms. SLCuV has a wide host range within the Cucurbitaceae, e.g., squash (Cucurbita pepo), watermelon and cucumber, and weeds, Chenopodium murale, Convolvulus spp., and Prosopis farcta (Ali-Shtayeh et al. 2014). Transmission SLCuV is transmitted by the whitefly, Bemisia tabaci. The virus can be acquired in as little as 30 min and transmitted with an inoculation access period of 30 min. There is a latent period of 48 h after acquisition before transmission can occur. Whiteflies can continue to transmit the virus for 7–10 days or perhaps for the life of the vector. There is no evidence for transovarial transmission of SLCuV (Cohen et al. 1983). This is consistent with a circulative, non-propagative mechanism of transmission (Rosell et al. 1999). SLCuV is not readily mechanically transmissible but can be transmissible by sapinoculation with difficulty to a limited number of hosts. Experimentally SLCuV has been introduced into Nicotiana benthamiana by Agrobacterium-mediated inoculation with cloned virus components (Lazarowitz and Lazdins 1991). Virion properties and genome SLCuV virions are geminated (twinned quasi-icosahedra) 22  38 nm (Cohen et al. 1983).

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The genome is a bipartite consisting of two circular, single-stranded DNA components. DNA-A contains 2634 nt (M38183 = NC_001936) and DNA-B of 2607 nt (M38182 = NC_001937) (Cohen et al. 1983; Lazarowitz and Lazdins 1991; Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the New World, the DNA-A component of SLCuV encodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for SLCuV.

Squash leaf curl Yunnan virus Taxonomic position Genus: Begomovirus

(SLCuYV)

Family: Geminiviridae

Geographical distribution SLCuYV infection in plants of Cucurbita pepo was reported from Yunnan, China (Xie and Zhou 2003). Symptoms and host(s) The virus-infected squash plants exhibit leaf curl and foliar yellow mosaic symptoms. The virus has only been identified in an unidentified squash (Cucurbita) spp. Transmission The transmission of the SLCuYV has not been investigated. It is likely that, in common with other begomoviruses, SLCuYV is transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. The virus is not mechanically sap-transmissible. Virion properties and genome The virion structure of SLCuYV has not been investigated. In common with all geminiviruses, the virions of SLCuYV are likely geminate (twinned quasi-icosahedra). With only two isolates of SLCuYV having been characterized, the precise nature of the genome (whether monopartite, monopartite betasatellite-associated, or bipartite) is unclear. The characterized genomes/genomic components of SLCuYV are each ~2714 nt (AJ420319 = NC_004651; KX388157) (Briddon 2001; Xie and Zhou 2003; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes/genomic components of SLCuYV encode the six genes typically encoded by monopartite begomoviruses and the DNA-A component of bipartite begomoviruses. The expression and function of the genes have not been investigated.

Squash mild leaf curl virus Taxonomic position Genus: Begomovirus

(SMLCuV)

Family: Geminiviridae

Geographical distribution SMLCuV infection in plants of Cucurbita pepo was reported from the Southwestern USA and Northern Mexico (Brown et al. 2002).

Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow)

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Symptoms and host(s) The virus-infected squash plants exhibit leaf curling and mild mottling symptoms. Transmission The transmission of the SMLCuV has not been investigated. It is likely that, in common with other begomoviruses, SMLCuV is transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Virion properties and genome The virion structure of SMLCuV has not been investigated. In common with all geminiviruses, the virions of SMLCuV are likely geminate (twinned quasi-icosahedra). SMLCuV is typical of the majority of begomoviruses native to the New World with a bipartite genome consisting of two circular, single-stranded DNA components. DNA-A contains 2612 nt (AF421552 = NC_004645) and DNA-B of 2578 nt (AF421553 = NC_004646) (Briddon 2001; Brown et al. 2002, 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the New World, the DNA-A component of SMLCuVencodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for DaChMV.

Squash mosaic virus

(SqMV)

Synonyms Cucurbit ring mosaic virus; Muskmelon mosaic virus; Pumpkin mosaic virus Taxonomic position Genus: Comovirus

Family: Secoviridae

Geographical distribution SqMV is widely distributed in plants of Cucurbita pepo grown in areas of China, the Czech Republic, Japan, Spain, Turkey, and the USA (Lockhart et al. 1982; CABI/EPPO 1997; Langham et al. 1997; Dikova 1998; Jossey and Babadoost 2008; Sevik and Toksoz 2008; Li et al. 2015). Symptoms and host(s) The virus-infected summer squash plants exhibit mosaic mottling, reduction of the leaves, and distortion symptoms. The fruits from infected plants are reduced in size and are mottled and malformed. This virus infects a number of cucurbits and exhibits various degrees of systemic symptoms. The range of symptoms expressed depends on the strain of SqMV and the host species and variety grown. SqMV host range in natural conditions is mainly limited to cucurbits, but the virus can also infect several Chenopodium species. Transmission The virus is transmitted by a number of beetle vectors, viz., Acalymma trivittata, A. thiemei, Diabrotica undecimpunctata, D. bivittula, Epilachna chrysomelina, and E. paenulata, in a non-persistent manner (Langham et al. 1997). These vectors acquire the virus within 5 min and can retain it for about 20 days. SqMV does not multiply in the vector, but it can be recovered from regurgitation of fluid, feces, and hemolymph. The virus is transmissible by mechanical sap-inoculation. The virus is seed-transmitted in number of cucurbits, at up to 35% in squash (Cucurbita pepo) and 10% in Cucumis melo. The virus is not

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Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow)

transmitted by pollen (Alvarez and Campbell 1978; Nolan and Campbell 1984; Lestari and Nurhayati 2014). Virion properties and genome The virions are isometric, non-enveloped of two types but similar in size 25–30 nm in diameter, and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome consists of two molecules of positivesense ssRNA designated as RNA-1 and RNA-2, which are encapsidated separately with two capsid proteins. RNA-1 is 5865 nt (AB054688 = NC_003799; EU421059) and contains a single long open reading frame encoding a polyprotein (2K, 58K helicase, VPg, 24K protease, 87K RNA-dependent RNA polymerase). RNA-2 is 3354 nt (AB054689 = NC_003800; KP223324, EU421060) and encodes two large over lapping polyproteins, from which finally four proteins are formed 52K RNA-2 replication protein, 38K larger coat protein and 21K smaller coat protein, and 42K movement protein (Haudenshield and Palukaitis 1998; Han et al. 2002; Hu et al. 2009; Bruening and Lomonossoff 2011; Li et al. 2015; Sanfacon 2015; Thompson et al. 2017).

Squash vein yellowing virus Taxonomic position Genus: Ipomovirus

(SqVYV)

Family: Potyviridae

Geographical distribution SqVYV infection in plants of Cucurbita pepo was reported from Florida and California (Adkins et al. 2007, 2008; Baker et al. 2008; Batuman et al. 2015). Symptoms and host(s) The virus-infected plants exhibit symptoms of vein yellowing and drying of leaves leading to death of the plants. The virus naturally infects watermelon and bitter gourd. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a non-persistent manner. At least 30 whiteflies for plant are needed for consistent transmission; and an acquisition access period of 1–2 h duration is required. The virus is also mechanically sap-transmissible to cucurbitaceous members (Adkins et al. 2008; Webb et al. 2006, 2012). Virion properties and genome The virions are non-enveloped, flexuous filaments measuring 840 nm long and 12–15 nm in diameter. The genome consists of a single molecule of positive-sense ssRNA of 9836 nt (EU259611 = NC_010521) which contains a single large open reading frame encoding a single polyprotein of 3172 amino acids (Li et al. 2008; Colinet 2011; Webster and Adkins 2012; Wylie et al. 2017).

Tobacco necrosis virus D Taxonomic position Genus: Betanecrovirus

(TNV-D)

Family: Tombusviridae

Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow)

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TNV-D was reported to infect glasshouse grown Cucurbita pepo plants in Northern Italy (Roggero and Lisa 1995). The virus-infected summer squash plants exhibit yellow spots on young leaves and necrotic symptoms on older leaves, petioles, and stems. The virus is transmitted by the fungal vector, Olpidium brassicae, and also by mechanical sap-inoculation. For more details of TNV-D, refer to Nicotiana tabacum.

C Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Cucurbita pepo was reported from Illinois, USA (Sammons and Barnett 1987; Jossey and Babadoost 2006). The virus-infected summer squash plants exhibit mild mosaic with leaf yellowing symptoms and also chlorotic spots. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV was identified in plants of Cucurbita pepo (Senator) showing severe mosaic and epinasty symptoms in Florida, USA (Padmanabhan et al. 2014). The virus is transmitted by the thrips vectors, is present in/on pollen, entering in to the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato black ring virus Taxonomic position Genus: Nepovirus

(TBRV)

Family: Secoviridae

TBRV infection in plants of Cucurbita pepo was reported from Poland (Pospieszny and Borodynko 2005). The virus-infected summer squash plants exhibit symptoms of mosaic, yellowing, stunting, leaf chlorosis, and fruit deformations. The virus is transmitted by the nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TBRV, refer to Solanum lycopersicum.

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

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Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow)

ToLCNDV infection in plants of Cucurbita pepo was reported from India, Southern Italy, and Spain (Tiwari et al. 2011; Juarez et al. 2014; Mnari-Hattab et al. 2015; Luigi et al. 2016; Panno et al. 2016; Anwar 2017). The virus-infected summer squash plants exhibit curling, vein swelling, and severe mosaic in young leaves, short internodes, and fruit skin roughness. The virus is transmitted by a whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is graft-transmissible. The virus failed to be transmissible through mechanical sap-inoculation, and seed. For more details of ToLCNDV, refer to Solanum lycopersicum.

Tomato leaf curl Palampur virus Taxonomic position Genus: Begomovirus

(ToLCPalV)

Family: Geminiviridae

ToLCPalV infection in plants of Cucurbita pepo was reported from India and Iran (Tiwari et al. 2010; Heydarnejad et al. 2013). The virus-infected summer squash plants exhibit mosaic, mottling, leaf curling, yellowing, and stunting symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is graft-transmissible. The virus failed to be transmissible through mechanical sap-inoculation, and seed. For more details of ToLCPalV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Cucurbita pepo was reported from North America and also reported from South Carolina, Chile, Denmark, Japan, New Zealand, Sweden, the UK, and Yugoslavia (Provvidenti 1986; Sammons et al. 1989). The virus-infected summer squash plants exhibit yellow mosaic, reduction in leaf size, shorter internodes, proliferation of the flower buds, and prominent ringspots on discolored fruits. The virus is transmitted by nematode vectors in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and also by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of butternut squash (Cucurbita moschata) was reported from Zimbabwe (Karavina et al. 2016; Karavina and Gubba 2017). The virus-infected plants exhibit symptoms of leaf mottling, crinkling, and mosaics. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow)

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

755

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Cucurbita pepo was reported from Cuba (Zubiaur et al. 2004). The virusinfected summer squash plants exhibit symptoms of curling and light yellowing of leaves. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not transmissible by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

Watermelon chlorotic stunt virus Taxonomic position Genus: Begomovirus

(WmCSV)

Family: Geminiviridae

WmCSV infection in plants of Cucurbita pepo was reported from Lebanon (Samsatly et al. 2012). The virus-infected summer squash plants exhibit leaf curling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by mechanical inoculation, but may be transmitted by grafting. For more details of WmCSV, refer to Citrullus lanatus.

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV infection in plants of Cucurbita pepo was reported from the Mediterranean area, China, Poland, Bosnia, Serbia, Yugoslavia, Libya, India, Argentina, and Herzegovina (Bhargava and Joshi 1960; Bhargava 1977; Al-Musa and Mansour 1982; Thouvenel et al. 1986; Sammons et al. 1989; Pearson and Liayanage 1997; Dukic et al. 2002; Krstic et al. 2002; Jossey and Babadoost 2008; Borodynko et al. 2009; Tiwari and Gao 2014; Trkulja et al. 2014; Svoboda et al. 2015; Perotto et al. 2016; Niu et al. 2017). The virus-infected summer squash plants exhibit mild to severe greening mosaic blistering, vein-banding, and distortion symptoms on leaves. The virus is transmitted by aphid vectors in a nonpersistent manner, and also by mechanical sap-inoculation. For more details of WMV, refer to Citrullus lanatus.

Zucchini green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(ZGMMV)

Family: Virgaviridae

Geographical distribution ZGMMV infection in plants of Cucurbita pepo was reported from Korea (Ryu et al. 2000).

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Symptoms and host(s) The virus-infected zucchini squash plants exhibit symptoms of mottling and severe mosaic on leaves and abnormal fruits. Under high temperature and humidity, symptoms are generally masked. Transmission No insect vector is known for this virus. The virus is transmissible through mechanical sap-inoculation, and also by contact between plants. Virion properties and genome The virions are 18 nm in diameter and have a predominant length of 300–310 nm. The genome consists of a single molecule of positive-sense ssRNA, which is 6513 nt (AJ295949 = NC_003878) encoding four proteins: of 131 kDa replication protein, 189 kDa readthrough replication protein, 28 kDa movement protein, and 17 kDa coat protein (Ryu et al. 2000; Yoon et al. 2002). The 30 -terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G50 pppG) (Zaitlin 2011; Adams et al. 2017).

Zucchini lethal chlorosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(ZLCV)

Family: Tospoviridae

Geographical distribution ZLCV infection in plants of Cucurbita pepo was reported from Brazil (Pozzer et al. 1996; Resende et al. 1997; Nakahara and Monteiro 1999; Bezerra et al. 1999). Symptoms and host(s) The virus-infected zucchini plants show severe mosaic, leaf distortion, stunting, and often death (Bezerra et al. 1999; Nakahara and Monteiro 1999). The virus infects several cucurbits under field conditions: Cucurbita pepo var. Caserta, C. maxima var. Alice, C. moschata var. Menina Brasileira, C. maxima  C. moschata hybrid Takayama, Citrullus lanatus var. Crimson Sweet, Cucumis sativus var. Safira, C. sativus var. Hokushin, C. anguria, Datura stramonium, and Luffa aegyptiaca (Giampan et al. 2007; Camelo-Garcia et al. 2015). Transmission The virus is transmitted by the thrips vector, Frankliniella zucchini, in a persistent-propagative manner (Nakahara and Monteiro 1999). The primary spread of the disease is significantly correlated with the population of thrips (Moreira et al. 2014). The virus is mechanically sap-transmissible. Virion properties and genome The virions are spherical and membrane bound, of 80–120 nm in diameter. The genome consists of three segments of ssRNA. The complete genome sequence is not available; however, partial genome sequence contains nucleocapsid protein (NP) (AF067069), movement protein (AF213676), and glycoprotein precursor (Gn/Gc) (AB274027) gene sequences which are available (Bezerra et al. 1999; Nagata et al. 2007). The virus is identified as a new tospovirus mainly based on the NP sequence (260 amino acids) that shares only 20–80% sequence identity with the other tospoviruses (Lima et al. 2016).

Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow)

Zucchini shoestring virus Taxonomic position Genus: Potyvirus

757

(ZSSV)

Family: Potyviridae

Geographical distribution ZSSV infection in plants of Cucurbita pepo was reported from South Africa (Ibaba et al. 2016). Symptoms and host(s) The virus-infected zucchini plants exhibit severe leaf filiformy and fruit deformation symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically saptransmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments measuring 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 10,308 nt excluding the poly(A) tail (KU355553) (Revers and Garcia 2015; Ibaba et al. 2016; Wylie et al. 2017).

Zucchini tigre mosaic virus Taxonomic position Genus: Potyvirus

(ZTMV)

Family: Potyviridae

Geographical distribution ZTMV was detected in plants of Cucurbita pepo from Guadeloupe (an overseas region of France), France, Florida (USA), and Venezuela (Romay et al. 2014a; Webb et al. 2016; Xiao et al. 2016; Abdalla and Ali 2017). Symptoms and host(s) The virus-infected zucchini plants exhibit characteristic mosaic symptoms. The virus naturally infects several cucurbits (Romay et al. 2014a). Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically saptransmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome consists of a single molecule of positive-sense ssRNA of 10346 nt (KC345607 = NC_023175) (Romay et al. 2014a).

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Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow)

Zucchini yellow fleck virus Taxonomic position Genus: Potyvirus

(ZYFV)

Family: Potyviridae

Geographical distribution ZYFV infection was first reported in plants of Cucurbita pepo from Southern Italy (Vovlas et al. 1981). The virus spreads in Italy, Iran, and Mediterranean countries (Desbiez et al. 2007; Safaeezadeh 2007). Symptoms and host(s) The virus-infected zucchini plants exhibit symptoms of yellow pinpoints, and systemic flecks on leaves and plants later become completely yellow, necrosis, stunting, and fruitlessness under severe infection. Transmission The virus is transmitted by the aphid vector, Myzus persicae, in a non-persistent manner. The virus is also transmissible by mechanical sap-inoculation to three to nine families; experimentally infected plants mostly show yellow local lesions and systemic mottling. Virion properties and genome The virions are non-enveloped, flexuous filaments measuring 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of c.9.5–10 kb. A partial genome sequence of 1711 nt is available (DQ641510) (Desbiez et al. 2007; Revers and Garcia 2015; Wylie et al. 2017).

Zucchini yellow mosaic virus Taxonomic position Genus: Potyvirus

(ZYMV)

Family: Potyviridae

Geographical distribution ZYMV infection was first reported in plants of Cucurbita pepo from Italy by Lisa et al. (1981). The virus spreads in Algeria, Asia, Iran, Iraq, Greece, Australia, Egypt, France, Germany, Israel, Italy, Jordan, Lebanon, Mauritius, Morocco, Serbia, Spain, Taiwan, Turkey, Czechoslovakia, Turkey, the UK, Oceania, Yugoslavia, Libya, Trinidad, Hungary, and South and North America (Adlerz et al. 1983; Davis and Yilmaz 1984; Wickizer et al. 1986; Greber et al. 1987, 1988; Crosslin et al. 1988; Ghorbani 1988; Chod and Jokes 1991; Kyriakopoulou and Varveri 1991; Desbiez and Lecoq 1997; Mahgoub et al. 1997; Gracia 2000; Basky et al. 2001; Bostan et al. 2002; Dukic et al. 2002; Krstic et al. 2002; CABI/EPPO 2003; Singh et al. 2003; Muller et al. 2006; Lecoq and Desbiez 2012; Sharma et al. 2013; Romay et al. 2014b; Tiwari and Gao 2014; Svoboda et al. 2015; Al-Kuwaiti et al. 2016; Chinnaraja et al. 2016). Three strains of ZYMV from distinct geographic origins were reported by Desbiez et al. (1996, 2002). Symptoms and host(s) The virus-infected zucchini plants develop symptoms of vein-clearing, mosaic, yellowing, leaf distortion, and severe filiformism. Fruits are generally severely misshapen with prominent knobs.

Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow)

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The virus has a narrow host range. In natural conditions, it infects mostly cultivated or wild cucurbits but also a few ornamental species (Althea, Begonia, and Delphinium) or weeds (Lecoq et al. 1981; Lisa et al. 1981; Lecoq and Desbiez 2008). Transmission The virus is transmitted by at least 26 aphid species in a non-persistent manner. A. craccivora, A. gossypii, Macrosiphum euphorbiae, and M. persicae are efficient vectors for this virus. Some aphid species (Lipaphis erysimi, Myzus ascalonicus) were not able to transmit this virus (Lecoq et al. 1981; Lisa et al. 1981; Adlerz 1987; Blua and Perring 1992; Yuan and Ullman 1996; Katis et al. 2006; Simmons et al. 2008; Vucurovic et al. 2012). The virus is transmissible by mechanical sap-inoculation to 11 families of dicotyledons. The virus is seed-transmitted in Cucurbita pepo at rates of 6.2–26.67% (Lecoq et al. 1981; Dodds et al. 1984; Davis and Mizuki 1986; Provvidenti and Robinson 1987; Schrijnwerkers et al. 1991; Tobias et al. 2008; Simmons et al. 2011; Lestari and Nurhayati 2014; Sevik and Balkaya 2015). Virion properties and genome The virions are non-enveloped, flexuous filaments with a clear modal length of 750 nm and 11 nm wide (Gal-On 2007). The genome consists of a single molecule of positive-sense ssRNA of 9591 nt (AF127929 = NC_003224) encoding a single polyprotein of 3080 amino acids, which matures into 10 different proteins (Purcifull et al. 1984; Davis 1986; Balint et al. 1990; Lin et al. 2001; Zhao et al. 2003; Hosseini et al. 2007; Neha et al. 2011; Azarfar et al. 2012; Novakova et al. 2014; Wylie et al. 2017).

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Moreira AS, Cônsoli RCM, Barbosa JC, Filho AB, Rezende JAM (2014) Spatiotemporal dynamics of Zucchini squash lethal chlorosis and its relationship with the Frankliniella zucchini population. J Phytopathol 162:481–490 Muller C, Bröther H, von Bargen S, Büttner C (2006) Zucchini yellow mosaic virus incidence and sources of virus infection in field-grown cucumbers and pumpkins in the Spreewald, Germany. J Plant Dis Prot 113:252–258 Nagata T, Carvalho KR, Sodré Rde A, Dutra LS, Oliveira PA, Noronha EF, Lovato FA, Resende Rde O, De Avila AC, Inoue-Nagata AK (2007) The glycoprotein gene of Chrysanthemum stem necrosis virus and Zucchini lethal chlorosis virus and molecular relationship with other tospoviruses. Virus Genes 35:785–793 Nakahara S, Monteiro RC (1999) Frankliniella zucchini (Thysanoptera: Thripidae), a new species and vector of Tospovirus in Brazil. Proc Entomol Soc Wash 101:290–294 Neha S, Vrat BS, Kumud J, Thakur PD, Rajinder K, Anil H (2011) Comparative in silico analysis of partial coat protein gene sequence of Zucchini yellow mosaic virus infecting summer Squash (Cucurbita pepo L.) isolated from India. J Proteomics Bioinform 4:068–073 Niu Y, Pang X, Cheng Y, Wang D, Guo S, Liu Y (2017) Molecular detection of watermelon mosaic virus associated with a serious mosaic disease on Cucurbita pepo L. in Shanxi, China. Virus Dis 28:315–319 Noa-Carrazana J, Silva-Rosales L (2001) First report of a Mexican isolate of Papaya mosaic virus in papaya (Carica papaya) and pumpkin (Cucurbita pepo). Plant Dis 85:558 Nolan PA, Campbell RN (1984) Squash mosaic virus detection in individual seeds and seed lots of cucurbits by enzymelinked immunosorbent assay. Plant Dis 68(11):971–975 Novakova S, Svoboda J, Glasa M (2014) Analysis of the complete sequences of two biologically distinct Zucchini yellow mosaic virus isolates further evidences the involvement of a single amino acid in the virus pathogenicity. Acta Virol 58:364–367 Omar AF, Bagdady NA (2012) Cucurbit aphid-borne yellows virus in Egypt. Phytoparasitica 40:177–184 Ozaslan M, Aytekin T, Bas B, HalilKilic I, DidemAfacan I, Dag DS (2006) Virus diseases of cucurbits in GaziantepTurkey. Plant Pathol J 5:24–27 Padmanabhan C, Gao S, Li R, Zhang S, Fei Z, Ling K-S (2014) Complete genome sequence of an emerging genotype of tobacco streak virus in the United States. https://doi.org/10.1128/genomeA.01138-14 Panno S, Iacono G, Davino M, Marchione S, Zappardo V, Bella P, Tomassoli L, Accotto GP, Davino S (2016) First report of tomato leaf curl New Delhi virus affecting zucchini squash in an important horticultural area of southern Italy. New Dis Rep 33:6 Pearson MN, Liayanage AS (1997) Records of cucurbit viruses infecting vegetable crops in Western Samoa. Australas Plant Pathol 28:188–191 Perotto MC, Celli MG, Pozzi EA, Luciani CE, Conci VC (2016) Occurrence and characterization of a severe isolate of Watermelon mosaic virus from Argentina. Eur J Plant Pathol 146:213–218 Polston JE, Hladky LL, Akad F, Wintermantel WM (2008) First report of Cucurbit yellow stunting disorder virus in cucurbits in Florida. Plant Dis 92:1251 Pospieszny H, Borodynko N (2005) First report of Tomato black ring virus (TBRV) in the natural infection of zucchini (Cucurbita pepo l. convar giromantiina) in Poland. J Plant Prot Res 45(4):321–325 Pozzer L, de Resende RO, Bezerra MI, Nagata T, Lima MI, Kitajima FW, de Avila AC (1996) Zucchini lethal chlorotic virus (ZLCV), a proposed new species in Tospovirus genus. Fitopatol Bras 21(Suppl):432 Provvidenti R (1986) Viral diseases of cucurbits and sources of resistance. ASPAC, Food & Fertilizer Technol. Center, Tech. Bulletin 93: 1–16 Provvidenti R, Robinson RW (1987) Lack of seed transmission in squash and melon plants infected with Zucchini yellow mosaic virus. Cucurbit Genet Coop Rep 10:81–82 Provvidenti R, Uyemoto JK (1973) Chlorotic leaf spotting of yellow summer squash caused by the severe strain of Bean yellow mosaic virus. Plant Dis Reptr 57:280–282 Purcifull DE, Adlerz WC, Simone GW, Hiebert E, Christie SR (1984) Serological relationships and partial characterization of Zucchini yellow mosaic virus isolated from squash in Florida. Plant Dis 68:230–233 Resende JAM, Galleti SR, Pozzer L, de Resende RO, de Avila AC, Scagliusi SM (1997) Incidence, biological and serological characteristics of a tospovirus infecting experimental fields of zucchini in Sao Paulo State, Brazil. Fitopatol Bras 22:92–95 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Revill PA, Ha CV, Porchun SC, Vu MT, Dale JL (2003) The complete nucleotide sequence of two distinct geminiviruses infecting cucurbits in Vietnam. Arch Virol 148(8):1523–1541 Rodriguez-Martinez D, Figueira AR, Duarte PSG, Galvino-Costa SBF, Olmedo JG (2015) First report and molecular characterization of an isolate of Papaya ringspot virus (PRSV-W) detected in pumpkin in Cuba. Biosci J 31:1133–1142 Roggero P, Lisa V (1995) Characterization of an isolate of tobacco necrosis virus from zucchini. J Phytopathol 143:485–489 Roggero P, Dellavalle G, Lisa V (1998) First report of Moroccan watermelon mosaic potyvirus in zucchini in Italy. Plant Dis 82:351

Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow)

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Romay G, Lecoq H, Desbiez C (2014a) Zucchini tigre mosaic virus is a distinct potyvirus in the papaya ringspot virus cluster: molecular and biological insights. Arch Virol 159(2):277–289 Romay G, Lecoq H, Desbiez C (2014b) Cucurbit crops and their viral diseases in Latin America and the Caribbean Islands: a review. J Plant Pathol 96(2):227–242 Rosell RC, Torres-Jerez I, Brown JK (1999) Tracing the geminivirus-whitefly transmission pathway by polymerase chain reaction in whitefly extracts, saliva, haemolymph, and honeydew. Phytopathology 89(3):239–246 Ryu KH, Min BE, Choi GS, Choi SH, Kwon SB, Noh GM, Yoon JY, Choi YM, Jang SH, Lee GP, Cho KH, Park WM (2000) Zucchini green mottle mosaic virus is a new tobamovirus; comparison of its coat protein gene with that of kyuri green mottle mosaic virus. Arch Virol 145:2325–2333 Safaeezadeh M (2007) First report of Melon necrotic spot virus and Zucchini yellow fleck virus in cucurbits in Iran. J Plant Pathol 89:302 Sammons B, Barnett OW (1987) Tobacco ring spot virus from squash grown in South Carolina and transmission of the virus through seed of smooth pigweed. Plant Dis 71:530–532 Sammons B, Barnett OW, Davis RF, Mizuki MK (1989) A survey of viruses infecting yellow summer squash in South Carolina. Plant Dis 73:401–404 Samsatly J, Sobh H, Jawhari M, Najjar C, Haidar A, Abou-Jawdah Y (2012) First report of Watermelon chlorotic stunt virus in Cucurbits in Lebanon. Plant Dis 96:1703 Sanfacon H (2015) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. In: Encyclopedia of Life Sciences. John Wiley and Sons Ltd, Chichester. https://doi.org/10.1002/9780470015902. a0000764.pub3. http://www.els.net Sanfacon H, Wellink J, Le Gall O, Karasev A, van der Vlugt R, Wetzel T (2009) Secoviridae: a proposed family of plant viruses within the order Picornavirales that combines the families Sequiviridae and Comoviridae, the unassigned genera Cheravirus and Sadwavirus, and the proposed genus Torradovirus. Arch Virol 154(5):899–907 Saritha RK, Bag TK, Loganathan M, Rai AB, Rai M (2011) First report of Squash leaf curl china virus causing mosaic symptoms on summer squash (Cucurbita pepo) grown in Varanasi district of India. Arch Phytopathol Plant Protect 44:179–185 Schrijnwerkers CCFM, Huijberts N, Bos L (1991) Zucchini yellow mosaic virus: two outbreaks in the Netherlands and seed transmissibility. Neth J Plant Pathol 97:187–191 Sevik MA, Balkaya A (2015) Seed transmissibility of viruses in winter squash landraces collected from the Black Sea region of Turkey. Plant Protect Sci 51:195–199 Sevik MA, Toksoz Y (2008) Occurrence of Squash mosaic virus (SqMV) infecting pumpkin and squash in Samsun, Turkey. J Turk Phytopathol 37:15–25 Sharma YR, Chohan JS (1973) Transmission of cucumis viruses 1 and 3 through seeds of cucurbits. Indian Phytopathol 26:596–598 Sharma M, Thakur PD, Gupta D, Thakur AK (2013) Identification of viruses and screening of summer squash (Cucurbita pepo) germplasm against viral disease under controlled conditions. Indian J Agric Sci 83:426–430 Simmons HE, Holmes EC, Stephenson AG (2008) Rapid evolutionary dynamics of Zucchini yellow mosaic virus. J Gen Virol 89:1081–1085 Simmons HE, Holmes EC, Gildow E, Bothe-Goralczyk MA, Stephenson AG (2011) Experimental verification of seed transmission of Zucchini yellow mosaic virus. Plant Dis 95:751–754 Singh SJ, Verma R, Ahlawat YS, Singh RK, Prakash S, Pant RP (2003) Natural occurrence of a yellow mosaic disease on zucchini in India caused by a potyvirus. Indian Phytopathol 56:174–179 Singh R, Raj SK, Prasad V (2008) Molecular characterization of a strain of Squash leaf curl China virus from North India. J Phytopathol 156:222–228 Sobh H, Samsatly J, Jawhari M, Najjar C, Haidar A, Abou-Jawdah Y (2012) First report of Squash leaf curl virus in cucurbits in Lebanon. Plant Dis 96(8):1231 Sufrin-Ringwald T, Lapidot M (2011) Characterization of a synergistic interaction between two cucurbit-infecting begomoviruses: Squash leaf curl virus and Watermelon chlorotic stunt virus. Phytopathology 101:281–289 Sun SE, Wang JQ, Chen S, Zhang SB, Zhang DY, Liu Y (2018) First report of Capsicum chlorosis orthotospovirus infecting Zucchini (Cucurbita pepo) in China. Plant Dis 102(10):2049 Svoboda J, Leisova-Svobodova L, Lecoq H (2011) First report of Cucurbit aphid-borne yellows virus in Squash in the Czech Republic. Plant Dis 95:220 Svoboda J, Grimova L, Zouhar M, Rysanek P, Homa H (2015) First report of Cucurbit aphid-borne yellows virus, Watermelon mosaic virus and Zucchini yellow mosaic virus infecting zucchini plants in Libya. Plant Dis 99:558 Tahir M, Haider MS, Briddon RW (2010) First report of Squash leaf curl China virus in Pakistan. Aust Plant Dis Notes 5:21–24 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J GenVirol 98:529–531

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Cucurbita pepo (Summer squash, Zucchini, Vegetable marrow)

Thouvenel J-C, Fauquet C, Fargette D (1986) Occurrence of Watermelon mosaic virus 1 in Niger. Plant Dis 70:173 Tiwari AK, Gao GP (2014) Viruses infecting Cucurbita pepo: current status and management. In: Kharwar RN et al. (eds) Microbial diversity and biotechnology in food security. Springer, India, pp 357–371 Tiwari AK, Mall S, Khan MS, Snehi SK, Sharma PK, Rao GP, Raj SK (2010) Detection and identification of Tomato leaf curl Palampur virus infecting Cucurbita pepo in India. Guanaxi Agric Sci 41:1291–1.295 Tiwari AK, Snehi SK, Singh R, Raj SK, Rao GP, Sharma PK (2011) Molecular identification and genetic diversity among six Begomovirus isolates affecting cultivation of cucurbitaceous crops in Uttar Pradesh, India. Arch Phytopathol Plant Protect 45:62–72 Tobias I, Szabo B, Salanki K, Sari L, Kuhlmann H, Palkovics L, Pitrat M (2008) Seedborne transmission of Zucchini yellow mosaic virus and Cucumber mosaic virus in Styrian Hulless group of Cucurbita pepo. Cucurbitaceae. (Proc) IXth EUCARPIA meeting on genetic and breeding of Cucurbitaceae. Avignon, pp 189–200 Trkulja V, Stojčic J, Kovačic D, Stankovic I, Vučurovic A, Bulajic A, Krstic B (2014) First report of Watermelon mosaic virus in zucchini squash in Bosnia and Herzegovina. Plant Dis 98:573 Tsai WS, Shih L, Green SK, Jan FJ (2007) Occurrence and molecular characterization of Squash leaf curl Philippines virus in Taiwan. Plant Dis 91(7):907 van der Vlugt RAA, Verbeek M, Dullemans AM, Wintermantel WM, Cuellar WJ, Fox A, Thompson JR (2015) Torradoviruses. Annu Rev Phytopathol 53:485–512 Verbeek M, van Bekkum PJ, Dullemans AM, van der Vlugt RA (2014) Torradoviruses are transmitted in a semi-persistent and stylet-borne manner by three whitefly vectors. Virus Res 186:55–60 Vovlas C, Hiebert E, Russo M (1981) Zucchini yellow fleck virus, a new potyvirus of Zucchini squash. Phytopathol Mediterr 20:123–128 Vucurovic A, Bulajic A, Stankovic I, Ristic D, Berenji J, Krstic B (2011) Characterization of Cucumber mosaic virus originating from cucurbits in Serbia. Pestic Phytomed 26(4):325–336 Vucurovic A, Bulajic A, Stankovic I, Ristic D, Berenji J, Jovic J, Krstic B (2012) Non-persistently aphid-borne viruses infecting pumpkin and squash in Serbia and partial characterization of Zucchini yellow mosaic virus isolates. Eur J Plant Pathol 133:935–947 Webb SE, Adkins S, Baker CA (2006) Whitefly transmission of a new virus infecting cucurbits in Florida. In: Holmes GJ (ed) Proceedings of Cucurbitaceae 2006. Universal Press, Raleigh, pp 309–316 Webb SE, Adkins S, Reitz SR (2012) Semi-persistent whitefly transmission of Squash vein yellowing virus causal agent of viral watermelon vine decline. Plant Dis 96:839–844 Webb SE, Badillo-Vargas IE, Purcifull DE, Hiebert E, Baker CA, Funderburk JE, Adkins S (2016) Zucchini tigre mosaic virus infection of cucurbits in Florida. Plant Dis 100:2540 Webster CG, Adkins S (2012) Low genetic diversity of Squash vein yellowing virus in wild and cultivated cucurbits in the U.S. suggests a recent introduction. Virus Res 163:520–527 Wickizer SL, Scott HA, McGuire JM (1986) Zucchini yellow mosaic virus in squash in Arkansas. Plant Dis 70:78 Wintermantel WM (2004) Pumpkin (Cucurbita maxima and C. pepo), a new host of Beet pseudo yellows virus in California. Plant Dis 88:82 Wong S, Thio SS, Shintaku MH, Palukaitis P (1999) The rate of cell-to-cell movement in squash of Cucumber mosaic virus is affected by sequences of the capsid protein. Mol Plant Microbe Interact 12:628–632 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J GenVirol 98:352–354 Xiang HY, Shang QX, Han CG, Li DW, Yu JL (2008) First report on the occurrence of Cucurbit aphid-borne yellows virus on nine cucurbitaceous species in China. Plant Pathol 57:390 Xiao L, Li YY, Tan GL, Lan PX, Zhong L, Liu Y, Li R, Li F (2016) First report of Zucchini tigre mosaic virus infecting several cucurbit plants in China. Plant Dis 100:1253 Xie Y, Zhou XP (2003) Molecular characterization of Squash leaf curl Yunnan virus, a new begomovirus and evidence for recombination. Arch Virol 148:2047–2054 Yakoubi S, Desbiez C, Fakhfakh H, Wipf-Scheibel C, Marrakchi M, Lecoq H (2007) Occurrence of Cucurbit yellow stunting disorder virus and Cucumber vein yellowing virus in Tunisia. J Plant Pathol 89(3):417–420 Yakoubi S, Lecoq H, Desbiez C (2008) Algerian watermelon mosaic virus (AWMV): a new potyvirus species in the PRSV cluster. Virus Genes 37:103–109 Yardimci N, Ozgonen H (2007) First report of Cucurbit aphid-borne yellows virus in Turkey. Aust Plant Dis Notes 2:59 Yoon JY, Min BE, Choi JK, Ryu KH (2002) Genome structure and production of biologically active in vitro transcripts of cucurbit-infecting Zucchini green mottle mosaic virus. Phytopathology 92:156–163 Yuan C, Ullman DE (1996) Comparison of efficiency and propensity as measures of vector importance in Zucchini yellow mosaic potyvirus transmission by Aphis gossypii and A. craccivora. Phytopathology 86:698–703 Zaitlin M (2011) Tobamovirus. In: Tidona C, Darai G (eds) The Springer index of viruses. Springer, New York. https://doi. org/10.1007/978-0-387-95919-1 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Cucurbita pepo var. styriaca (Oil seed pumpkin)

767

Zhang YJ, Li GF, Li MF (2009) Occurrence of Cucumber green mottle mosaic virus on cucurbitaceous plants in China. Plant Dis 93:200 Zhao MF, Chen J, Zheng HY, Adams MJ, Chen JP (2003) Molecular analysis of Zucchini yellow mosaic virus isolates from Hangzhou. China J Phytopathol 151:307–311 Zindovic J, Manglli A, Hrncic S, Radonjic S, Perovic T, Tomassoli L (2017) First report of Cucurbit aphid-borne yellows virus affecting summer squash and melon in Montenegro. J Plant Pathol 99:299 Zubiaur YM, Fonseca D, Quinones M, Palenzuela I (2004) Presence of Tomato yellow leaf curl virus infecting squash (Cucurbita pepo) in Cuba. Aust Plant Dis Notes 88:572

Cucurbita pepo var. styriaca (Oil seed pumpkin) Family: Cucurbitaceae

Oilseed

Cucumber mosaic virus

(CMV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV infection in plants of Cucurbita pepo var. styriaca was reported from Serbia (Vucurovic et al. 2009, 2011b). The virus-infected oilseed pumpkin plants exhibit mosaic symptoms on the leaves and dark green streaks on the stem and petioles. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(CABYV)

Family: Luteoviridae

CABYV infection in plants of Cucurbita pepo “Olinka” was reported from Serbia (Vucurovic et al. 2011a). The virus-infected oilseed pumpkin plants exhibit symptoms of severe yellowing and thickening of older leaves. The virus is transmitted by aphid vectors, Myzus persicae, M. euphorbiae, and Aphis gossypii, in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of CABYV, refer to Cucumis melo.

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV infection in plants of Cucurbita pepo var. styriaca was reported from Serbia (Vucurovic et al. 2009). The virus-infected oilseed pumpkin plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical in plants of sap-inoculation. For more details of WMV, refer to Citrullus lanatus.

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Cudrania tricuspidata (Chinese mulberry)

Zucchini yellow mosaic virus Taxonomic position Genus: Potyvirus

(ZYMV)

Family: Potyviridae

ZYMV infection in plants of Cucurbita pepo var. styriaca was reported from Serbia (Vucurovic et al. 2009). The virus-infected oilseed pumpkin plants show blister-like symptoms, and fruits display bump-like bulges. The virus is transmittable in a non-persistent manner by a number of aphid species to the plant, and also through the seed (0.3–15.3%) (Riedle-Bauer et al. 2002; Tobias and Palkovics 2003). The virus is mechanically sap-transmissible. For more details of ZYMV, refer to Cucurbita pepo.

References Riedle-Bauer M, Saurez M, Reinprecht HJ (2002) Seed transmission and natural reservoirs of Zucchini yellow mosaic virus in Cucurbita pepo var. styriaca. J Plant Dis Protect 109:200–206 Tobias I, Palkovics L (2003) Characterization of Hungarian isolates of zucchini yellow mosaic virus (ZYMV, potyvirus) transmitted by seeds of Cucurbita pepo var. Styriaca. Pest Manag Sci 59:493–497 Vucurovic A, Bulajic A, Dekic I, Ristic D, Berenji J, Krstic B (2009) Presence and distribution of oilseed pumpkin viruses and molecular detection of Zucchini yellow mosaic virus. Pestic Fitomed 24:85–94 Vucurovic A, Bulajic A, Stankovic I, Ristic D, Berenji J, Jovic J, Krstic B (2011a) First report of the occurrence of cucurbit aphid-borne yellows virus on oilseed pumpkin in Serbia. Plant Dis 95:1035 Vucurovic A, Bulajic A, Stankovic I, Ristic D, Berenji J, Krstic B (2011b) Characterization of Cucumber mosaic virus originating from cucurbits in Serbia. Pestic Phytomed 26(4):325–336

Cudrania tricuspidata (Chinese mulberry) Family: Moraceae

Medicinal

Bean common mosaic virus Taxonomic position Genus: Potyvirus

(BCMV)

Family: Potyviridae

BCMV infection in plants of Cudrania tricuspidata was reported from Korea (Seo et al. 2015). The virus-infected Chinese mulberry plants exhibit symptoms of mosaic, yellowing, and distortion on the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BCMV, refer to Phaseolus vulgaris.

References Seo J-K, Kang M, Shin OJ, Kwak H-R, Kim M-K, Choi H-S, Ko S-J (2015) First Report of Bean common mosaic virus in Cudrania tricuspidata in Korea. Plant Dis 99:292

Cupressus sempervirens (Graveyard cypress)

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Cullen australasicum (Cullen) Synonyms Psoralea australasica Family: Fabaceae

C Forage plant

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

(AMV)

Family: Bromoviridae

AMV infection in plants of Cullen australasicum was reported from South Australia (Nair et al. 2009). The virus-infected Cullen plants exhibit severe yellow mosaic and leaf distortion symptoms. The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sapinoculation. For more details of AMV, refer to Medicago sativa.

References Nair RM, Habili N, Randles JW (2009) Infection of Cullen australasicum (syn. Psoralea australasica) with Alfalfa mosaic virus. Aust Plant Dis Notes 4:46–48

Cupressus sempervirens (Graveyard cypress) Family: Cupressaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Cupressus sempervirens was reported from Iran (Ghotbi and Sharaeen 2012). The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

770

Curcuma spp.

TSWV infection in plants of Cupressus sempervirens was reported from Iran (Ghotbi and Sharaeen 2012). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Reference Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses affecting propagated ornamentals in Northern Iran. Int Res Microbiol 3:373–381

Curcuma spp. Family: Zingiberaceae

Spice crop

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Curcuma spp. was reported from the Czech Republic (Mertelik et al. 2000). The virus-infected curcuma plants exhibit symptoms of chlorotic spots and patterns. The virus is transmitted by a thrips vector, Frankliniella occidentalis, in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. Propagation of rhizomes from the infected plants is another important factor for virus spread. For more details of INSV, refer to Impatiens spp.

Reference Mertelik J, Mokra V, Gotzova B, Gabrielova S (2000) First report of impatiens necrotic spot virus in the Czech Republic. Plant Dis 84:1045

Cuscuta spp. Family: Convolvulaceae

Weed host

Arabis mosaic virus

(ArMV)

Taxonomic position Genus: Nepovirus

Family: Secoviridae

ArMV infection in plants of Cuscuta spp. was reported from Iran (Ghotbi and Shahraeen 2005). The virusinfected cuscuta plants exhibit mosaic, leaf chlorosis, small necrotic lesions, leaf malformation, and

Cuscuta spp.

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deformation symptoms. This virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in plants of Cuscuta campestris was reported from Bulgaria (Dikova 2006). The virusinfected cuscuta plants exhibit symptoms of yellow-orange-colored stems and fruits. The virus is transmitted by nematode vectors and also by mechanical sap-inoculation to a large number of host plants. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Cuscuta spp. was reported from Spain and Cyprus (Jorda et al. 2001; Papayiannis et al. 2011). The virus-infected cuscuta plants exhibit symptomless infections. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

Tomato yellow ring virus

(TYRV)

Synonyms Tomato Varamin virus (ToVV) Taxonomic position TYRV is a tentative member of the genus Orthotospovirus and family Tospoviridae ToVV infection in plants of Cuscuta spp. was reported from Iran (Ghotbi et al. 2005). The virusinfected cuscuta plants may or may not exhibit any conspicuous virus symptoms. The virus is transmitted by a thrips vector, Thrips tabaci, in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of TYRV, refer to Solanum lycopersicum.

References Dikova B (2006) Establishment of tobacco rattle virus (trv) in weeds and Cuscuta. Biotechnol Biotechnol Equip 20:42–48 Ghotbi T, Shahraeen N (2005) First report on incidence of Arabis mosaic virus (ArMV, Nepovirus) on ornamental plants in Iran. Iranian J Plant Pathol 41(2):305–306 Ghotbi T, Shahraeen N, Winter S (2005) Occurrence of tospoviruses in ornamental and weed species in Markazi and Tehran provinces in Iran. Plant Dis 89(4):425–429 Jorda C, Font I, Martinez P, Juarez M, Ortega A, Lacasa A (2001) Current status and new natural hosts of tomato yellow leaf curl virus (TYLCV) in Spain. Plant Dis 85:445 Papayiannis L, Katis NI, Idris AM, Brown JK (2011) Identification of weed hosts of Tomato yellow leaf curl virus in Cyprus. Plant Dis 95:120–125

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Cyamopsis tetragonoloba (Guar/Cluster Bean)

Cyamopsis tetragonoloba (Guar/Cluster Bean) Family: Fabaceae

Grain legume

Bean common mosaic virus

(BCMV)

Synonyms Guar green sterile virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

BCMV infection in plants of Cyamopsis tetragonoloba was reported from Georgia and South Africa and also occurs wherever this crop is grown (Ben-Moshe et al. 1991; Rey et al. 1995; Gillaspie et al. 1998). The virus-infected guar plants exhibit symptoms of mosaic, small necrotic lesions on the leaves, and reduced inflorescences. The virus is transmitted by several aphid vectors, including Acyrthosiphon pisum, Aphis craccivora, A. fabae, Myzus persicae, and other spp. in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation to a wide range of hosts. The virus is seed-transmitted through the seed of Cyamopsis tetragonoloba up to 94% (Gillaspie et al. 1998). For more details of BCMV, refer to Phaseolus vulgaris.

Papaya leaf curl virus Genus: Begomovirus

(PaLCuV) Family: Geminiviridae

PaLCuV infection in plants of Cyamopsis tetragonoloba was reported from Pakistan (Tahir et al. 2017). The virus-infected cluster bean plants exhibit leaf curl symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. The virus is not mechanically sap-transmissible. PaLCuV was shown to be associated with Tomato leaf curl betasatellite. Additionally, some cluster bean plants were shown to also harbour Cotton leaf curl Multan alphasatellite (Tahir et al. 2017). For more details of PaLCuV, refer to Carica papaya.

Tobacco necrosis virus

(TNV)

Taxonomic position TNV is a tentative member of the genus Necrovirus and family Tombusviridae TNV infection in plants of Cyamopsis tetragonolobus was reported from India (Ramachandraiah et al. 1979). The virus-infected guar plants exhibit symptoms of elongated, light-colored, necrotic areas with dark margins and were stunted, twisted, and necrotic streaks on the petals. The virus is transmitted by the zoospores of the fungus Olpidium brassicae, and the virus is also mechanically sap-transmissible. For more details of TNV, refer to Nicotiana tabacum.

Cyamopsis tetragonoloba (Guar/Cluster Bean)

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

773

(TRSV)

Family: Secoviridae

TRSV infection in plants of Cyamopsis tetragonoloba was reported from the United States (Orellana 1966). The virus-infected guar plants exhibit symptoms of top stem necrosis and leaf symptoms consisting of circular to subcircular necrotic lesions of light brown with thin darker margins. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sapinoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Cyamopsis tetragonoloba was reported from India (Bhat et al. 2002; Sivaprasad et al. 2012). The virus-infected guar plants exhibit symptoms of foliar mosaic and necrotic spotting as well as necrotic streaks on buds and stems. The virus is transmitted by thrips vectors, the virus present in/on pollen, entering in to the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Cyamopsis tetragonoloba was reported from Pakistan (Zaidi et al. 2017). The virus-infected guar plants exhibit leaf yellowing and curling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not transmissible by mechanical sap-inoculation. For more details of TYLCV, refer to Solanum lycopersicum.

References Ben-Moshe H, Rey MEC, Sibara M, Garnett HM, Beck B (1991) A new green-sterile disease of guar in South Africa. Plant Dis 75:638–643 Bhat AI, Jain RK, Ramiah M (2002) Detection of Tobacco streak virus from sunflower and other crops by reverse transcription polymerase chain reaction. Indian Phytopathol 55:216–218 Gillaspie AG Jr, Pappu HR, Jain RK, Rey MEC, Hopkins MS, Pinnow DL, Morris JB (1998) Characteristics of a latent potyvirus seed-borne in guar and of guar greensterile virus. Plant Dis 82:765–770 Orellana RG (1966) A new occurrence of tobacco ringspot of guar in the United States. Plant Dis Reptr 50:7–10 Ramachandraiah M, Venkatarathnam P, Sulochana CB (1979) Tobacco necrosis virus: occurrence in India. Plant Dis Reptr 63:949–951

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Cycas spp.

Rey MEC, Smith A, Johnson S, Khitsane S (1995) Preliminary characterization of a potyvirus, the causal agent of greensterile disease of guar. Plant Dis 79:172–176 Sivaprasad Y, Reddy BVB, Sujitha A, Sai Gopal DVR (2012) First report of tobacco streak virus infecting Cyamopsis tetragonoloba. J Plant Pathol 94:S4.96 Tahir MN, Mansoor S, Briddon RW, Amin I (2017) Begomovirus and associated satellite components infecting cluster bean (Cyamopsis tetragonoloba) in Pakistan. J Phytopathol 165(2):115–122 Zaidi SS, Shakir S, Farooq M, Amin I, Mansoor S (2017) First report of a novel strain of tomato yellow leaf curl virus causing yellow leaf curl disease on cluster bean in Pakistan. Plant Dis 101:1071

Cycas spp. Family: Cycadaceae

Ornamental

Cycas necrotic stunt virus Taxonomic position Genus: Nepovirus

(CNSV)

Family: Secoviridae

Geographical distribution CNSV was first reported in plants of Cycas revoluta from Chiba Prefecture, Japan, by Kusunoki et al. (1975). The virus spreads in Japan (Kusunoki et al. 1986). Symptoms and host(s) The virus-infected cycas plants exhibit symptoms of dwarfing and twisting of young leaves and chlorotic or necrotic spots on mature leaves. The diseased plants declined in growth year after year, and severely affected plants were finally killed. Transmission The virus is mechanically sap-transmissible and the host range of the virus is limited. Out of 39 species in 12 families, only 10 species in Aizoaceae, Amaranthaceae, and Cycadaceae were susceptible to the virus. On some Chenopodium species, chlorotic local lesions and systemic mottling were produced. The virus was transmitted through seeds from disease plants of C. amaranticolor and C. serotinum at the rate of 29% and 82%, respectively (Kusunoki et al. 1986). Virion properties and genome The virions are isometric, non-enveloped of two types but similar in size 25–30 nm in diameter, and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is bipartite, linear positive-sense, single-stranded RNA. RNA1 consists of 7471 nt (AB073147 = NC_003791) and RNA2 of 4667 nt (AB073148 = NC_003792) (Hanada et al. 1986; Han et al. 2002; Sanfacon et al. 2009; Dunez and Le Gall 2011; Sanfacon 2015; Thompson et al. 2017).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(NSV)

Cycas spp.

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INSV was reported to infecting plants of Cycas spp. in Iran (Ghotbi and Shahraeen 2012; Ghotbi 2013). The virus-infected cycas plants exhibit symptoms including necrotic leaf spot, chlorosis, and stunting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

IYSV infection in plants of Cycas spp. was reported from Markazi and Tehran provinces in Iran (Shahraeen and Ghotbi 2003; Ghotbi et al. 2005). The virus is transmitted by onion thrips (Thrips tabaci) in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of IYSV, refer to Iris spp. or Allium cepa.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV was reported to infecting plants of Cycas spp. in Iran (Ghotbi and Shahraeen 2012). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Dunez J, Le Gall O (2011) Nepovirus. Comoviridae. The Springer index of Viruses. Springer, New York, pp 361–369. https://doi.org/10.1007/978-0-387-95919-1_55. Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iranian J Plant Pathol 49:41 Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses affecting propagated ornamentals in Northern Iran. Int Res J Microbiol 3:373–381 Ghotbi T, Shahraeen N, Winter S (2005) Occurrence of Tospoviruses in ornamental and weed species in Markazi and Tehran provinces in Iran. Plant Dis 89:425–429 Han SS, Karasev AV, Ieki H, Iwanami T (2002) Nucleotide sequence and taxonomy of Cycas necrotic stunt virus. Arch Virol 147:2207–2214 Hanada K, Kusunoki M, Iwaki M (1986) Properties of virus particles, nucleic acid and coat protein of cycas necrotic stunt virus. Phytopathol Soc Jpn 52:422–427 Kusunoki M, Nagai Y, Yamashita S, Doi Y, Yora K (1975) Small spherical virus identified in Cycas necrotic stunt plants. Ann Phytopathol Soc Jpn 41:285 Kusunoki M, Hanada K, Iwaki M, Chang MU, Doi Y, Yora K (1986) Cycas necrotic stunt virus, a new member of nepoviruses found in Cycas revoluta; host range, purification, serology and some properties. Ann Phytopathol Soc Jpn 52:302–311 Sanfacon H (2015) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. In: eLS. Wiley, Chichester. http://www.els.net https://doi.org/10.1002/9780470015902.a0000764.pub3 Sanfacon H, Wellink J, Le Gall O, Karasev A, van der Vlugt R, Wetzel T (2009) Secoviridae: a proposed family of plant viruses within the order Picornavirales that combines the families Sequiviridae and Comoviridae, the unassigned genera Cheravirus and Sadwavirus, and the proposed genus Torradovirus. Arch Virol 154(5):899–907

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Cyclamen persicum (Cyclamen)

Shahraeen N, Ghotbi T (2003) Natural occurrence of different Tospovirus species infecting ornamentals and other agricultural crops in Iran. Abstract 23.26 of papers offered at the 8th International Congress of Plant Pathology, Christchurch, New Zealand, 2–7 February 2003 (p 307) Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J Gen Virol 98:529–531

Cyclamen persicum (Cyclamen) Family: Primulaceae

Ornamental

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Cyclamen persicum was reported from the Netherlands, Taiwan, and Italy (Loebenstein et al. 1995; Chen 2003). The virus-infected cyclamen plants exhibit streaking and malformation symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Cyclamen persicum was reported from Pennsylvania (USA), Japan, New Zealand, Hungary, the USA, and Italy (Vaira et al. 1992a, b; Daughtrey et al. 1997; Goto and Natsuaki 2001; Goto et al. 2001; Toth et al. 2007; Elliott et al. 2009). The virus-infected cyclamen plants exhibit different leaf symptoms such as mottling, necrosis, ringspots, chlorosis, distortion, and stunted growth. Lesions initiated at the base of the leaf blade may radiate along several veins causing an oak leaf pattern. Chlorotic leaf lesions develop into necrotic spots or concentric rings. Leaf collapse can result from petiole necrosis or coalescence of numerous ringspots. These symptoms can vary with the stage of growth of the host and with a variety of cultural conditions. The virus is transmitted by a thrips vector, Frankliniella occidentalis, in a persistent-propagative manner, and also by mechanical sapinoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Potato virus X

(PVX)

Taxonomic position Genus: Potexvirus

Family: Alphaflexiviridae

Cyclamen persicum (Cyclamen)

777

PVX infection in plants of Cyclamen persicum has been reported sporadically from Denmark (Paludan 1965). The virus-infected cyclamen plants exhibit symptoms of flower distortion and necrotic streaking of petals. The virus is transmitted by mechanical sap-inoculation, and also by contact between plants. For more details of PVX, refer to Solanum tuberosum.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

C

(TMV)

Family: Virgaviridae

TMV infection in plants of Cyclamen persicum has been recorded in the Netherlands (Wegman 1966). The virus-infected cyclamen plants exhibit mild symptoms of chlorosis and stunting of plants. There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Cyclamen persicum was reported from the USA and Canada (Allen and Matteoni 1988; Hausbeck et al. 1992; Daughtrey 1996; Jones and Baker 1990). On cyclamen the most distinctive symptom of TSWV is yellow ringspots and whitish blotches, spots, and streaks on the leaves. Brown streaks, in addition to rings, may be present on the petioles. Flowers are often dwarfed and malformed. Corms may be elongated, and, when cut, the vascular tissues appear as brown streaks. Affected plants stop growing, and brown spots appear on leaves and leaf edges, leading to death of the plants (Vicchi et al. 1992). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Allen WR, Matteoni JA (1988) Cyclamen ring spot: epidemics in Ontario greenhouses caused by the Tomato spotted wilt virus. Can J Pathol 10:41–46 Chen YK (2003) Occurrence of cucumber mosaic virus in ornamental plants and perspectives of transgenic control. Ph.D. Thesis, Wageningen University, The Netherlands. p 144 Daughtrey ML (1996) Detection and identification of tospoviruses in greenhouses. Acta Hortic 431:90–98 Daughtrey ML, Jones RK, Moyer JW, Daub ME, Baker JR (1997) Tospoviruses strike the greenhouse industry: INSV has become a major pathogen on flower crops. Plant Dis 81(11):1220–1230 Elliott DR, Lebas BSM, Ochoa-Corona FM, Tang J, Alexander BJR (2009) Investigation of impatiens necrotic spot virus outbreaks in New Zealand. Aust Plant Pathol 38:490–495 Goto T, Natsuaki T (2001) Occurrence of necrotic spot of cyclamen caused by Impatiens necrotic spot virus (INSV) (in Japanese). Plant Protect 55:541–544 Goto T, Sazarashi H, Nozawa H, Nakayama K, Natsuaki T (2001) Necrotic spot disease of cyclamen (Cyclamen persicum) and exacum (Exacum affine) caused by impatiens necrotic spot virus (INSV) (in Japanese with English summary). Annu Rep Kanto-Tosan Plant Prot Soc 48:97–100

778

Cyclanthera pedata (Kaywa)

Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among greenhouse ornamentals in Pennsylvania. Plant Dis 76:795–800 Jones RK, Baker R (1990) TSWV: symptoms, host range and spread. Proceedings of a USDA Workshop, “Virus-thripsplant interactions of Tomato spotted wilt virus”, Beltsville, Maryland, p 89–93 Loebenstein G, Lawson RH, Brunt AA (1995) Virus and virus-like diseases of bulb and flowering crops. Wiley, UK, p 543 Paludan N (1965) Virussygdome hos syclamen. Maanesdovers 419:83–88 Toth EK, Kriston E, Takacs A, Bajtek M, Kazinczi G, Horvath J (2007) First report of impatiens necrotic spot virus in ornamental plants in Hungary. Plant Dis 91:331 Vaira AM, Lisa V, Dellavelle G (1992a) Distribution of tospovirus in Liguria, Italy. 5th Int Plant virus epidemiology symposium, “Viruses, vectors and the environment”, valenzano Bari p 23–24 Vaira AM, Gallo S, Lisa V (1992b) Nuove infezioni da due Tospovirus (tomato spotted wilt e impatiens necrotic spot) in Liguria. Informatore Fitopatologco 42:37–42 Vicchi V, Bellardi MG, Bertaccini A (1992) Le infezione da TSWV dilagano in Italia: nuovi casi in altertre regioni. L’Inf Agrario 23:53–56 Wegman J (1966) Jaarverslag 1965, Proefstation voor de Bloemisterij in Nederland te Aalsmeer. 134 pp

Cyclanthera pedata (Kaywa) Family: Cucurbitaceae

Vegetable

Papaya ringspot virus Taxonomic position Genus: Potyvirus

(PRSV)

Family: Potyviridae

PRSV pathotype W infection in plants of Cyclanthera pedata was reported from Brazil (Rezende 2000). The virus-infected Kaywa plants exhibit intense mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PRSV, refer to Carica papaya.

Reference Rezende JAM (2000) Cyclanthera pedata var. edulis: new host of papaya ringspot virus-type W in Brazil. Plant Dis 84:1155

Cydonia oblonga (Quince) Family: Rosaceae

Edible fruit

Apple chlorotic leaf spot virus Taxonomic position Genus: Trichovirus

(ACLSV)

Family: Betaflexiviridae

Cydonia oblonga (Quince)

779

ACLSV infection in plants of Cydonia oblonga was reported from the USA, England, Canada, New Zealand, South Africa, Greece, Iran, Turkey, and Europe (Cropley et al. 1963; Fridlund 1986; Mathioudakis et al. 2007; Akbas and Ilhan 2008; Rana et al. 2008; Birisik and Baloglu 2010; Alemzadeh et al. 2016). The virus-infected quince plants exhibit irregularly shaped yellow spots and curl. Shoots grow slowly and die, depending on the degree of plant susceptibility. In some cultivars pale green, yellow ring and line patterns occur on the leaves. No natural vector has been reported for this virus. The virus is transmissible by mechanical sapinoculation to a number of herbaceous hosts. The use of infected clonal root stocks and use of infected trees as source of propagation materials are the major source of virus spread. The virus is transmissible by grafting. The virus is not transmitted by seed or pollen. For more details of ACLSV, refer to Malus domestica.

Apple mosaic virus

(ApMV)

Taxonomic position Genus: Ilarvirus

Family: Bromoviridae

ApMV infection in plants of Cydonia oblonga was reported from Turkey (Dursunoglu and Ertunc 2008). No insect vector is known for this virus. The virus is mechanically sap-transmissible and by grafting. For more details of ApMV, refer to Malus domestica.

Apple stem grooving virus Taxonomic position Genus: Capillovirus

(ASGV)

Family: Betaflexiviridae

ASGV infection in plants of Cydonia oblonga was reported from Turkey and India (Birisik and Baloglu 2010; Negi et al. 2010). The virus-infected quince plants exhibit symptoms of mosaic, leaf deformation, and fruit malformation. No vector has been identified for this virus. The virus is mechanically saptransmissible. The virus is transmissible by grafting and disseminated by infected propagative material. For more details of ASGV, refer to Malus domestica.

Apple stem pitting virus Taxonomic position Genus: Foveavirus

(ASPV)

Family: Betaflexiviridae

ASPV infection in plants of Cydonia oblonga was reported from Turkey and Greece (Mathioudakis et al. 2006, 2012; Birisik and Baloglu 2010; Ilbagi et al. 2013). The virus-infected quince plants exhibit symptoms of mosaic, leaf deformation, and fruit malformation. The virus is transmitted by means not involving a vector. The virus is transmissible by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants. The use of bud sticks from infected plants is the major mode of spread of this disease. For more details of ASPV, refer to Malus domestica.

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Cymbidium spp.

Pear blister canker viroid Taxonomic position Genus: Apscaviroid

(PBCVd)

Family: Pospiviroidae

PBCVd infection in plants of Cydonia oblonga was reported from Australia (Joyce et al. 2006). The viroid-infected quince plants do not exhibit characteristic symptoms. The viroid is transmissible by grafting, budding, and mechanical sap-inoculation. For more details of PBCVd, refer to Pyrus communis.

References Akbas B, Ilhan D (2008) First report of apple chlorotic leaf spot virus in quince (Cydonia oblonga mill.) in Turkey. Acta Hortic 781:161–166 Alemzadeh E, Katsiani AT, Efthimiou K, Katis NI (2016) Occurrence of apple chlorotic leaf spot virus in apple and quince in Southern Iran. J Plant Pathol 98:177 Birisik N, Baloglu S (2010). Evaluation of the presence and symptomology of viruses in commercial quince orchards in Turkey. In: 21st international conference on virus and other graft transmissible diseases of fruit crops. Julius-KühnArchiv, 427:257 Cropley R, Wolfwinkel LD, Posnette AF (1963) The identification of some viruses infecting apple, pear and quince. Phytopathol Mediterr 2:132–136 Dursunoglu S, Ertunc F (2008) Distribution of apple mosaic ilarvirus (ApMV) in Turkey. Acta Hortic 781:131–134 Fridlund PR (1986) Symptom intensities in pear and quince cultivars incited by the virus that causes pear ring pattern mosaic and apple chlorotic leaf spot. Zast Bilja 37:327–333 Ilbagi H, Katsiani A, Uzunogullari N, Katis N, Citir A (2013) First report of apple stem pitting virus on pear and quince orchards in northern Turkey. J Plant Pathol 95:452 Joyce PA, Constable FE, Crosslin J, Eastwell K, Howell WE, Rodoni BC (2006) Characterization of pear blister canker viroid isolates from Australian pome fruit orchards. Australas Plant Pathol 35:465–471 Mathioudakis MM, Maliogka VI, Dovas CI, Vasilakakis M, Katis NI (2006) First record of the apple stem pitting virus (ASPV) in quince in Greece. J Plant Pathol 88:225 Mathioudakis MM, Candresse T, Katis NI (2007) First report of apple chlorotic leaf spot virus in quince in Greece. Plant Dis 91:462 Mathioudakis MM, Candresse T, Barone M, Ragozzino A, Katis NI (2012) Cydonia japonica, Pyrus calleryana and P. amygdaliformis: three new ornamental or wild hosts of apple stem pitting virus. Virus Genes 44:319–322 Negi A, Rana T, Kumar Y, Ram R, Hallan V, Zaidi AA (2010) Analysis of the coat protein gene of Indian strain of Apple stem grooving virus. J Plant Biochem Biotechnol 19:91–94 Rana T, Chandel V, Hallan V, Zaidi AA (2008) Cydonia oblonga as reservoir of apple chlorotic leaf spot virus in India. Plant Pathol 57:393

Cymbidium spp. Family: Orchidaceae

Ornamental

Calanthe mild mosaic virus Synonyms Cymbidium potyvirus

(CalMMV)

Cymbidium spp.

Taxonomic position Genus: Potyvirus

781

Family: Potyviridae

CalMMV infection in plants of Cymbidium pendulum and C. tigrinum was reported from India (Singh et al. 2007). The virus-infected cymbidium plants exhibit mild mosaic and stunting symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CalMMV, refer to Calanthe spp.

Cymbidium chlorotic mosaic virus Taxonomic position Genus: Sobemovirus

(CyCMV)

Family: Solemoviridae

Geographical distribution CyCMV infection in plants of Cymbidium goeringii and C. forrestii was reported from Japan (Kondo et al. 1994, 1996, 2015) and in C. tortisepalum in China (Zhang et al. 2015). Symptoms and host(s) The virus-infected cymbidium plants exhibit symptoms of stunting and chlorotic streaks on newly developed leaves; in China necrotic spotting was reported in C. tortisepalum. Transmission The virus is mechanically transmissible only to Cymbidium orchids. Virion properties and genome The virions are isometric, non-enveloped, and 28 nm in diameter. The genome is polycistronic, positive-sense, single-stranded RNA (ssRNA) of 4083 nt (LC019764 = NC_027123). The genome consists of two overlapping ORFs, ORF2a, and ORF2b (Kondo et al. 2015).

Cymbidium mosaic virus Taxonomic position Genus: Potexvirus

(CymMV)

Family: Alphaflexiviridae

Geographical distribution CymMV infection in plants of Cymbidium spp. was first reported from California by Jensen (1950). The virus is worldwide in distribution (Jensen 1970; Zettler et al. 1990; Hu et al. 1993; Wong et al. 1994; Youn et al. 1997; Park et al. 1998; Ryu et al. 1998; Sherpa et al. 2003, 2007; Navalinskiene et al. 2005; Pant et al. 2010). Symptoms and host(s) The virus-infected Cymbidium cultivars exhibit symptoms such as chlorotic streaking. The virus also exhibits a mosaic of irregularly shaped chlorotic and/or necrotic lesions to appear on infected hosts. Additionally, infected plants will show smaller yields. Sometimes plants may display chlorotic rings while others will display symptoms in lesions. Sunken patches can also be observed on leaves, and flower breaking is commonly noticed. However, certain infected plants may be symptomless but still

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Cymbidium spp.

have virus enough to infect other neighboring plants. The virus infects a wide range of orchid genera but does not naturally infect plants other than orchids (Pearson and Cole 1986; Lawson 1990). Transmission The virus is mechanically sap-transmissible to Chenopodium amaranticolor and Datura stramonium. The virus is also transmitted by contact between infected and healthy plants. The virus also spreads through contact by contaminated cutting tools, equipment, human hands, potting media, or water (Allen 2012). No vector transmission is reported. The virus is not transmitted by seed but is transmitted through pollen (Hamilton and Valentine 1984). Virion properties and genome The virions are filamentous, flexuous rods with a modal length of c. 475 nm and 13 nm width. The genome consists of single-stranded positive-sense RNA of 6227 nt (U62963 = NC_001812) and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type, 18–27 kDa in size (Wong et al. 1997; Kim et al. 1998; Adams et al. 2004).

Cymbidium ringspot virus Taxonomic position Genus: Tombusvirus

(CymRSV)

Family: Tombusviridae

Geographical distribution CymRSV was first reported in plants of Cymbidium spp. from Sussex, UK, by Hollings and Stone (1963). The virus is cosmopolitan in distribution (Hollings 1972; Hollings et al. 1977). Symptoms and host(s) The virus-infected cymbidium plants exhibit chlorotic ring-mottling symptoms. Transmission The virus is transmitted by means not involving a vector. The virus is transmissible by mechanical sapinoculation to more than nine families. The virus is not transmitted by seed. Virion properties and genome The virions are isometric and icosahedral (T = 3), non-enveloped, and about 32–35 nm in diameter. The genome is a monopartite, linear, single-stranded, positive-sense RNA of 4733 nt (X15511 = NC_003532) (Grieco et al. 1989; White 2011).

Odontoglossum ringspot virus Taxonomic position Genus: Tobamovirus

(ORSV)

Family: Virgaviridae

ORSV infection in plants of Cymbidium spp. is distributed worldwide (Toussaint et al. 1984; Pearson and Cole 1986; Park et al. 1990a, b, 1998; Zettler et al. 1990; Ryu et al. 1998; Navalinskiene et al. 2005; Sherpa et al. 2006; Chen et al. 2010; Pant et al. 2010; Kumar et al. 2017). The virus-infected cymbidium

Cymbidium spp.

783

plants induce necrotic rings on the leaves and color breaking and distortion of flowers. This virus infection reduces plant vigor, growth rate, and flower quality. This virus is not transmitted by natural vectors but spread by contaminated tools, equipment, and potting media during division of plants and harvest of flowers. The virus is mechanically sap-transmissible and also by contact between plants. For more details of ORSV, refer to Odontoglossum grande.

C Orchid fleck dichorhavirus Taxonomic position Genus: Dichorhavirus

(OFV)

Family: Rhabdoviridae

Geographical distribution OFV was first reported in plants of Cymbidium spp. from Japan by Doi et al. (1969) and Chang et al. (1973). The virus spreads in Australia, Brazil, Denmark, Germany, Japan, Korea, New Zealand, Central America, China, and the USA (Kitajima et al. 1974; Lesemann and Doraiswamy 1975; Kondo et al. 1995, 2006; Freitas-Astua et al. 2002; Peng et al. 2017). Symptoms and host(s) The virus-infected cymbidium plants display chlorotic and necrotic ringspots on leaves (Kondo et al. 2006). Transmission The virus is transmitted by the mite vector, Brevipalpus californicus, in a persistent propagative manner, and is efficiently transmitted by both adults and nymphs, but not by larvae. Viruliferous mites retain their infectivity for 3 weeks on a virus-immune host. The virus is transmissible by mechanical sap-inoculation but is not transmitted by seed (Maeda et al. 1998; Kondo et al. 2003). Virion properties and genome The virions are bacilliform and non-enveloped particles of 32–35  100–110 nm. The genome comprises two molecules of negative-sense single-stranded RNA: RNA1 consists of 6413 nt (AB244417 = NC_009608) and RNA2 of 6001 nt (AB244418 = NC_009609). RNA1 contains five ORFs, and RNA2 has a single ORF encoding an RNA polymerase (Chang et al. 1976; Kondo et al. 2003, 2006, 2009; Peng et al. 2013; Dietzgen et al. 2014; Kondo et al. 2014; Walker et al. 2018).

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Cymbidium spp. was reported from the USA (Goff and Corbett 1977a). The virus-infected cymbidium plants exhibit faint chlorotic leaf streak symptoms. The virus is transmitted by nematode vectors in a non-persistent manner. The virus is transmissible by mechanical sapinoculation, and also by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

784

Cymbidium spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Cymbidium spp. was reported from the USA (Goff and Corbett 1977b). The virus-infected cymbidium plants exhibit faint chlorotic leaf streak symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Allen CD (2012) Transmission of Cymbidium mosaic virus in oncidium orchids by Periplaneta australasiae. MS Thesis submitted to University of Maryland Chang MU, Arai K, Doi Y, Yora K (1973) Short rod virus inducing necrotic spots on orchids – tentative orchid fleck virus. Ann Phytopath Soc Japan 39:171 Chang MU, Arai K, Doi Y, Yora K (1976) Morphology and intracellular appearance of orchid fleck virus. Ann Phytopath Soc Japan 42:156–157 Chen CC, Lin JY, Cheng YH, Chiang FL (2010) Distribution of Odontoglossum ringspot virus on plants of Cymbidium spp. and its application on virus detection. Plant Prot Bull 52:17–24 Dietzgen RG, Kuhn JH, Clawson AN, Freitas-Astua J, Goodin MM, Kitajima EW, Kondo H, Wetzel T, Whitfield AE (2014) Dichorhavirus: a proposed new genus for Brevipalpus mite-transmitted, nuclear, bacilliform, bipartite, negative-strand RNA plant viruses. Arch Virol 159:607–619 Doi Y, Arai K, Yora K (1969) Distribution of bacilliform virus particles in Masaki mosaic disease and cymbidium ring spot disease. Ann Phytopathol Soc Japan 35:388 Freitas-Astua J, Moreira L, Rivera C, Rodriguez CM, Kitajima EW (2002) First report of Orchid fleck vírus in Costa Rica. Plant Dis 86:1402 Goff LM, Corbett MK (1977a) Association of Tomato ring spot virus with a chlorotic leaf streak of Cymbidium orchids. Phytopathology 67:1096–1100 Goff LM, Corbett MK (1977b) Association of Tomato spotted wilt virus with a chlorotic leaf streak of Cymbidium orchids. Phytopathology 67:1096–1100 Grieco F, Burgyan J, Russo M (1989) The nucleotide sequence of cymbidium ringspot virus RNA. Nucleic Acids Res 17:6383 Hamilton RI, Valentine B (1984) Infection of orchid pollen by Cymbidium mosaic and Odontoglossum ring spot viruses. Can J Plant Pathol 6:185–190 Hollings M (1972) Orchids-cymbidium ring spot. Glass House Crops Research Institute Annual Report. p 104 Hollings M, Stone OM (1963) Cymbidium ring spot (previously undescribed virus). Glasshouse Crops Research Institute Annual Report. p 89 Hollings M, Stone OM, Barton RJ (1977) Pathology, soil transmission and characterization of cymbidium ringspot, a virus from cymbidium orchids and white clover (Trifolium repens). Ann Appl Biol 85:233–248 Hu JS, Ferreira S, Wang M, Xu MQ (1993) Detection of cymbidium mosaic virus, odontoglossum ringspot virus, tomato spotted wilt virus, and potyviruses infecting orchids in Hawaii. Plant Dis 77:464–468 Jensen DD (1950) Mosaic of cymbidium orchids. Phytopathology 40:966–967 Jensen DD (1970) Virus diseases of orchids in the Netherlands. Neth J Plant Pathol 76:135–139 Kim JD, Koo KY, Chang MU (1998) Genome characterization of a Korean isolate of cymbidium mosaic virus. Mol Cells 8:181–188 Kitajima EW, Blumenschein A, Costa AS (1974) Rodlike particles associated with ringspot symptoms in several orchid species in Brazil. Phytopathol Z 81:280–286 Kondo H, Maeda T, Inouye N (1994) A new sobemovirus isolated from Cymbidium goeringii: Cymbidium chlorotic mosaic virus. Japan Plant Pathol Manabu 60:396–397 Kondo H, Matsumoto J, Maeda T, Inouye N (1995) Host range and some properties of orchid fleck virus isolated from oriental Cymbidium in Japan. Bull Res Inst Bioresour Okayama Univ 3:151–161

Cymbidium spp.

785

Kondo H, Maeda T, Mitsuhata K, Inouye N (1996) Detection of the viruses occurring in oriental Cymbidium in Japan. Bull Res Inst Bioresources Okayama Univ 4:149–162 Kondo H, Maeda T, Tamada T (2003) Orchid fleck virus: brevipalpus californicus mite 0transmission, biological properties and genome structure. Exp Appl Acarol 30:215–223 Kondo H, Takanori M, Shirako Y, Tamada T (2006) Orchid fleck virus is a rhabdovirus with an unusual bipartite genome. J Gen Virol 87:2413–2421 Kondo H, Maeda T, Tamada T (2009) Identification and characterization of structural proteins of orchid fleck virus. Arch Virol 154:37–45 Kondo H, Maruyama K, Chiba S, Andika IB, Suzuki N (2014) Transcriptional mapping of the messenger and leader RNAs of orchid fleck virus, a bisegmented negative-strand RNA virus. Virology 452–453:166–174 Kondo H, Takemoto S, Maruyama K, Chiba S, Andika IB, Suzuki N (2015) Cymbidium chlorotic mosaic virus, a new sobemovirus isolated from a spring orchid (Cymbidium goeringii) in Japan. Arch Virol 160:2099–2104 Kumar A, Jailani AAK, Roy A, Mandal B (2017) The occurrence, biology and genomic properties of tobamoviruses infecting crop plants in India. In: Mandal B, Rao GP, Baranwal VK, Jain RK (eds) A century of plant virology in India. Springer Nature, Singapore, pp 429–443. isbn:978-981-10-5671-0 Lawson RH (1990) Orchid viruses and their control. In: Pridgeon AM, Tillman LL (eds) Handbook on orchid pests and diseases, Rev. edn. American Orchid Society, West Palm Beach, pp 66–101 Lesemann D, Doraiswamy S (1975) Bullet-shaped virus-like particles in chlorotic and necrotic leaf lesions of orchids. Phytopathol Z 83:27–29 Maeda T, Kondo H, Mitsuhata K, Tamada T (1998) Evidence that orchid fleck virus is efficiently transmitted in a persistent manner by the mite Brevipalpus californicus. In: Proc of the 7th Int Congr Plant Pathology, Edinburgh, Scotland, 3:13–18 Navalinskiene M, Raugalas J, Samuitiene M (2005) Viral diseases of flower plants 16. Identification of viruses affecting orchids (Cymbidium Sw.). Biologija 2:29–34 Pant RP, Mrinal Das KB, Pun KB, Ramachandran P, Medhi RP (2010) Occurrence of cymbidium mosaic and Odontoglossum ringspot viruses in orchid germplasm of Sikkim and Darjeeling hills, their identification and diagnosis. Indian Phytopathol 63:326–332 Park WM, Yoon KE, Chung SY, Ryu KH (1990a) Purification and serological detection of Odontoglossum ringspot virus isolated from Cymbidium goeringii in Korea. Korean J Plant Pathol 6:474–481 Park WM, Kim WG, Ryu KH, Yoon KE (1990b) Detection of Odontoglossum ringspot virus and Cymbidium mosaic virus from cultivated orchids by immunosorbent electron microscopy. J Kor Soc Hortic Sci 31:417–422 Park WM, Shim KB, Kim SJ, Ryu KH (1998) Detection of Cymbidium mosaic virus and Odontoglossum ring spot virus by ELISA and RT-PCR from cultivated orchids in Korea. Korean J Plant Pathol 14:130–135 Pearson MN, Cole JS (1986) The effects of Cymbidium mosaic virus and Odontoglossum ringspot virus on growth of cymbidium orchids. J Phytopathol 117:193–197 Peng DW, Zheng GH, Zheng ZZ, Tong QX, Ming YL (2013) Orchid fleck virus: an unclassified bipartite, negative-sense RNA plant virus. Arch Virol 158:313–323 Peng DW, Zheng GH, Tong QX, Zheng ZZ, Ming YL (2017) First report of orchid fleck dichorhavirus from Cymbidium sp. in China. Plant Dis 101:514 Ryu KH, Kim SJ, Park WM (1998) Incidence of cymbidium mosaic virus and odontoglossum ringspot virus in cymbidium and other orchids in Korea. J Kor Soc Hortic Sci 39(2):213–217 Sherpa AR, Hallan V, Ram R, Vij SP, Pathak P, Garg ID, Zaidi AA (2003) First report of Cymbidium mosaic virus (CymMV) in orchids from India. Plant Pathol 52:788 Sherpa AR, Bag TK, Hallan V, Zaidi AA (2006) Detection of Odontoglossum ringspot virus in orchids from Sikkim, India. Aust Plant Pathol 35:69–71 Sherpa AR, Bag TK, Hallan V, Zaidi AA (2007) Incidence of cymbidium mosaic virus (CymMV) in Sikkim. Indian Phytopathol 60:133–136 Singh MK, Sherpa AR, Hallan V, Zaidi AA (2007) A potyvirus in Cymbidium spp. in northern India. Austral Plant Dis Notes 2:11–13 Toussaint A, Dekegel D, Vanheule G (1984) Distribution of odontoglossum ringspot virus in apical meristems of infected cymbidium cultivars. Physiol Plant Pathol 25:297–305 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Rhabdoviridae. J Gen Virol 99:447–448 White KA (2011) Tombusvirus. Tombusviridae. In: The Springer index of Viruses. Springer, New York, pp 1917–1926. https://doi.org/10.1007/978-0-387-95919-1_314 Wong SM, Chang CG, Lee YH, Tan K, Zettler FW (1994) Incidence of cymbidium mosaic and odontoglossum ringspot viruses and their significance in orchid culture in Singapore. Crop Prot 13:235–239

C

786

Cymbopogon citratus (Lemon grass)

Wong SM, Mahtani PH, Lee KC, Yu HH, Tan Y, Neo KK, Chan Y, Wu M, Chng CG (1997) Cymbidium mosaic potexvirus RNA: complete nucleotide sequence and phylogenetic analysis. Arch Virol 142:383–391 Youn HS, Kim JD, Koo YB, Ko SJ, Chang MU (1997) Coat protein gene and 30 - noncoding region of a new Korean isolate of Cymbidium mosaic virus. Mol Cells 7:34–39 Zettler FH, Ko NJ, Wisler GC, Eliott MS, Wong SM (1990) Viruses of orchids and their control. Plant Dis 74:621–626 Zhang Z, Zheng K, Dong J, McBeath JH, Wang X (2015) Characterization of Cymbidium chlorotic mosaic virus from Cymbidium tortisepalum in China. J Plant Pathol 97:535–538

Cymbopogon citratus (Lemon grass) Family: Poaceae

Commercial crop

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Cymbopogon citratus was reported from India (Raj et al. 2007). The virusinfected lemon grass plants exhibit symptoms of severe yellow stripes and tip necrosis of leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Reference Raj SK, Kumar S, Pratap D, Vishnoi R, Choudhari S, Chandra S (2007) Natural occurrence of Cucumber mosaic virus on lemongrass (Cymbopogon citratus), a new record. Australas Plant Dis 2(1):95–96

Cynanchum spp. (Keunjorong) Family: Apocynaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Medicinal

(CMV)

Family: Bromoviridae

CMV infection in plants of Cynanchum taiwanianum was reported from Taiwan (Yang 2009; Yang et al. 2014). The virus-infected Keunjorong plants show mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Cynanchum spp. (Keunjorong)

787

Keunjorong mosaic virus Taxonomic position Genus: Potyvirus

(KjMV)

Family: Potyviridae

Geographical distribution KjMV infection in plants of Cynanchum wilfordii was reported from Korea (Lee et al. 2010; Nam et al. 2013). Symptoms and host(s) The virus-infected Keunjorong plants exhibit symptoms of mosaic, mottle, necrosis, yellowing, chlorotic spot, and malformation. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is mechanically saptransmissible to several herbaceous hosts, but only Chenopodium quinoa developed symptoms (chlorotic local lesions on the inoculated leaves and systemic chlorotic spots on upper leaves). Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome is a single molecule of linear positive-sense, single-stranded RNA of 9626 nt (JF838187 = NC_016159) (Lee et al. 2010; Nam et al. 2013; Wylie et al. 2017).

Onion yellow dwarf virus Taxonomic position Genus: Potyvirus

(OYDV)

Family: Potyviridae

OYDV infection in plants of Cynanchum spp. was reported from Egypt (Abdel Wahab et al. 2009). The virus-infected Keunjorong plants exhibit symptoms of mosaic and yellow streak symptoms, striping, curling and distortion of flower stems, and reduction in the number of flowers and seeds. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of OYDV, refer to Allium cepa.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Cynanchum acutum was reported from Israel (Cohen et al. 1988). The virus-infected Keunjorong plants exhibit yellow leaf curl symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

C

788

Cynara cardunculus (Cardoon)

References Abdel Wahab AS, Elnagar S, El-Sheikh MAK (2009) Incidence of aphid-borne Onion yellow dwarf virus (OYDV) in alliaceae crops and associated weeds in Egypt. 4th Conference on Recent Technologies in Agriculture Cohen S, Kern J, Harpaz I, Ben-Joseph R (1988) Epidemiological studies of the tomato yellow leaf curl virus (TYLCV) in the Jordan Valley, Israel. Phytoparasitica 16:259–270 Lee JH, Park SJ, Nam M, Kim MJ, Lee JB, Sohn HR, Choi HS, Kim JS, Lee JS, Moon JS, Lee SH (2010) Identification of a new potyvirus, Keunjorong mosaic virus in Cynanchum wilfordii and C. auriculatum. Res Plant Dis 16(3):238–246 Nam M, Lee JH, Choi HS, Lim HS, Moon JS, Lee SH (2013) Complete genome sequence of keunjorong mosaic virus, a potyvirus from Cynanchum wilfordii. Arch Virol 158:1817–1820 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Yang SH (2009) Characterization of Cucumber mosaic virus isolated from Taiwan swallowwort (Cynanchum taiwanianum Yaent Protection National Pingtung University of Science and Technology Pingtung, 76 pp Yang SH, Chen TH, Tzeng J, Lee WL, Tsao LY, Chen CC (2014) Characterization of Cucumber mosaic virus isolated from Taiwan swallowwort (Cynanchum taiwanianum Yamazaki) in Taiwan. Plant Protect Bull (Taipei) 56(4):109–136

Cynara cardunculus (Cardoon) Family: Asteraceae

Leafy vegetable

Artichoke yellow ring spot virus Taxonomic position Genus: Nepovirus

(AYRSV)

Family: Secoviridae

AYRSV infection in plants of Cynara cardunculus was reported from Sicily (southern Italy) (Martelli and Rana 1976; Rana et al. 1978). The virus-infected cardoon plants exhibit symptoms of striking yellow ringspots and line patterns in the leaves. There is no known vector for this virus. The virus is mechanically sap-transmissible. For more details of AYRSV, refer to Cynara cardunculus var. scolymus.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Cynara cardunculus was first reported from Italy (Lisa 1971). The virus spreads in the France, southern Italy, Tunisia, Spain, and Slovenia (Lisa 1971). The virus-infected cardoon plants generally exhibit mottling symptoms, while certain cardoon cultivars are symptomless. The virus is transmitted by the aphid vector, Myzus persicae, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

Cynara cardunculus var. scolymus (Globe artichoke)

789

TMV infection in plants of Cynara cardunculus was reported from Italy (Lisa 1971). The virus-infected plants exhibit symptoms of faint yellowing of the leaves. There is no known vector for this virus. The virus is mechanically sap-transmissible and also through contact between plants. For more details of TMV, refer to Nicotiana tabacum.

C

References Lisa V (1971) Two viruses naturally infecting cultivated cardoon (Cynara cardunculus L.). Phytopathol Mediterr 10:231–237 Martelli GP, Rana GL (1976) Viruses and virus diseases of globe artichoke and cardoon. Nuovi studi sul Carciofo. Proceedings of the 2nd Congresso Internazionale sul Carciofo, Bari 1973. pp 811–830 Rana GL, Rosciglione B, Cannizzaro G (1978) La maculatura anulare gialla del cardo e del carciofo. Phytopathol Mediterr 17:63–64

Cynara cardunculus var. scolymus (Globe artichoke) Family: Asteraceae

Leafy vegetable

Artichoke Aegean ringspot virus Taxonomic position Genus: Nepovirus

(AARSV)

Family: Secoviridae

Geographical distribution AARSV infection in plants of Cynara cardunculus var. scolymus was reported from Turkey and Greece (Kyriakopoulou and Bem 1982; Murant et al. 1996; Gallitelli et al. 2004, 2012; Rana et al. 2005). Symptoms and host(s) The virus-infected globe artichoke plants show yellow blotches and mild mottling symptoms. Transmission There is no known vector for this virus (Roca et al. 1986). The virus is mechanically sap-transmissible and has moderately wide experimental host range, infecting 22 species out of 40 in 7 dicotyledonous families (Rana et al. 1985). Virion properties and genome The virions are isometric, non-enveloped of two types but similar in size, are 25–30 nm in diameter, and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is bipartite, linear positive-sense, single-stranded RNA (Sanfacon et al. 2009; Dunez and Le Gall 2011; Sanfacon 2015; Thompson et al. 2017) No sequence information is available.

Artichoke Italian latent virus Taxonomic position Genus: Nepovirus

(AILV)

Family: Secoviridae

790

Cynara cardunculus var. scolymus (Globe artichoke)

Geographical distribution AILV infection in plants of Cynara cardunculus var. scolymus was first reported from Apulia (Southern Italy) by Majorana and Rana (1970a). The virus also occurs in Greece, Bulgaria, and Italy (Martelli et al. 1981; Rana and Kyriakopoulou 1982; Gallitelli et al. 2004, 2012; Rana et al. 2005). AILV at northeast Peloponnesus (Greece) is also known as artichoke patchy chlorotic stunting disease (Kyriakopoulou 1995). Symptoms and host(s) At Apulia (Southern Italy), AILV was recovered from symptomless artichokes (Majorana and Rana 1970a). Occasionally this disease exhibits mild yellowing of leaves along with stunting of plants (Martelli and Rana 1976). The natural hosts of AILV include cultivated species of chicory, gladiolus, grapevine, lettuce, and pelargonium. Transmission The virus is transmitted by nematode vectors, Longidorus apulus and L. fasciatus, in a non-persistent manner (Roca et al. 1975, 1982; Kyriakopoulou 1995; Brown et al. 1997; Brown and Trudgill 1998). The virus is mechanically sap-transmissible and has a wide natural and experimental host ranges, including 63 species in 12 dicotyledonous families (Savino et al. 1977). The virus is seed-transmitted in artichoke up to 5%, where it was detected in both seed coats and fully expanded cotyledons (Bottalico et al. 2002). Virion properties and genome The virions are isometric, non-enveloped of two types but similar in size, are 25–30 nm in diameter, and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is bipartite, linear positive-sense, single-stranded RNA. The sequences of RNA1 (7338 nt, LT608395) and RNA2 (4630 nt, LT608396) of a grapevine isolate are available (Sanfacon et al. 2009; Dunez and Le Gall 2011; Sanfacon 2015; Elbeaino et al. 2017; Thompson et al. 2017).

Artichoke latent virus Taxonomic position Genus: Macluravirus

(ArLV)

Family: Potyviridae

Geographical distribution ArLV infection in plants of Cynara cardunculus var. scolymus was reported from the Mediterranean region, Morocco, France, Tunisia, Sardinia, Italy, from California (USA), and Brazil (Costa et al. 1959; Fischer and Lockhart 1974; Foddai et al. 1977, 1983; Baricevich et al. 1995; Gallitelli et al. 2004, 2012; Rana et al. 1982, 1992, 2005; Ciuffo et al. 2011; Erkan et al. 2014). Symptoms and host(s) This virus-infected globe artichoke plants are symptomless (Marrou and Mehani 1964). Transmission The virus is transmitted by aphid vectors, Myzus persicae, Brachycaudus cardui, and Aphis fabae in a non-persistent manner (Rana et al. 1982). The virus is mechanically sap-transmissible. The virus is seed-transmitted to the extent of 5–10% in artichoke (Bottalico et al. 2002).

Cynara cardunculus var. scolymus (Globe artichoke)

791

Virion properties and genome The virions are non-enveloped, flexuous filaments, 650–675 nm long, and 13–16 nm in diameter. The genome consists of a single molecule of linear positive-sense ssRNA of 8287 nt (KP405232 = NC_026759) (Lopez-Moya et al. 2009; Foster 2011; Minutillo et al. 2015; Wylie et al. 2017).

Artichoke latent virus M

(ArLVM)

Taxonomic position ArLVM is a tentative member of the genus Carlavirus and family Betaflexiviridae Geographical distribution ArLVM infection in plants of Cynara cardunculus var. scolymus was reported from Apulia (Italy), Europe, and the Mediterranean Basin (Di Franco et al. 1989; Gallitelli et al. 2004; Rana et al. 2005). Symptoms and host(s) This virus-infected globe artichoke plants are symptomless. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is not mechanically sap-transmissible. Virion properties and genome The virions are slightly flexuous filaments, 610–700 nm in length, and 12–15 nm in diameter. The genome consists of a single molecule of linear single-stranded RNA (King et al. 2012). No sequence is currently available.

Artichoke latent virus S

(ArLVS)

Taxonomic position ArLVS is a tentative member of the genus Carlavirus and family Betaflexiviridae Geographical distribution ArLVS infection in plants of Cynara cardunculus var. scolymus was reported from the USA, Italy, Morocco, Spain, and Brazil (Fischer and Lockhart 1974; Rana et al. 1989, 2005; Gallitelli et al. 2004; Ortega et al. 2005). Symptoms and host(s) This virus-infected globe artichoke plants are symptomless (Costa et al. 1959; Majorana and Rana 1970b). Transmission The virus is transmitted by the aphid vector, Myzus persicae, in a non-persistent manner. The virus is mechanically sap-transmissible to less than three families.

C

792

Cynara cardunculus var. scolymus (Globe artichoke)

Virion properties and genome The virions are flexuous filaments, with a clear modal length of 664 nm and 12 nm wide. The genome consists of single-stranded RNA (King et al. 2012). No sequence is currently available.

Artichoke mottled crinkle virus Taxonomic position Genus: Macluravirus

(AMCV)

Family: Tombusviridae

Geographical distribution AMCV infection in plants of Cynara cardunculus var. scolymus was reported from Italy, Morocco, Malta, Greece, and Tunisia (Martelli et al. 1976, 1981; Rana and Cherif 1981; Rana and Kyriakopoulou 1982; Gallitelli et al. 2004, 2012; Rana et al. 2005). No sequence is currently available. Symptoms and host(s) The virus-infected plants exhibit symptoms of severe deformation, mottling, and crinkling of the leaves. Growth and yield of symptomatic plants are severely affected, flower heads are distorted, and plants may die. If the plant survives, the new foliage develops poorly and often shows bright chrome yellow discoloration. Transmission The virus is soil-borne without the aid of a biological vector. The virus is transmissible by mechanical sap-inoculation, and also through infected planting material used for propagation. Virion properties and genome The virions are isometric and icosahedral (T = 3), non-enveloped, and about 32–35 nm in diameter. The genome is a monopartite, linear, positive-sense, single-stranded RNA of 4789 nt (X62493 = NC_001339) (Tavazza et al. 1994; White 2011).

Artichoke yellow ringspot virus Taxonomic position Genus: Nepovirus

(AYRSV)

Family: Secoviridae

Geographical distribution AYRSV infection in plants of Cynara cardunculus var. scolymus was reported from Italy, Greece, and Turkey (Kyriakopoulou and Bem 1973; Martelli and Rana 1976; Kyriakopoulou et al. 1985; Gallitelli et al. 2004, 2012; Rana et al. 2005; Paylan et al. 2013). Symptoms and host(s) The virus-infected globe artichoke plants exhibit bright yellow blotches, ringspots, and line patterns on the leaves, occasionally followed by extensive necrosis. The virus infects 36 plant species in 14 families, 6 of which are agricultural crops (artichoke, cardoon, tobacco, broad bean, French bean, and cucumber) (Rana et al. 1980).

Cynara cardunculus var. scolymus (Globe artichoke)

793

Transmission The patchy distribution of the diseased plants suggests that a soil-inhabiting vector, possibly a nematode may be involved. The virus is mechanically sap-transmissible, and experimental host range is wide, including 56 species in dicotyledonous families (Rana et al. 1980). The virus is easily transmitted by seed (Kyriakopoulou et al. 1985) and vegetative propagation. Virion properties and genome The virions are isometric, non-enveloped of two types but similar in size, and 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is bipartite, linear positive-sense, single-stranded RNA. A partial sequence of 5873 nt is available (AM087671) (Sanfacon et al. 2009; Dunez and Le Gall 2011; Sanfacon 2015; Thompson et al. 2017).

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Cynara cardunculus var. scolymus was reported from Apulia (Italy) and Greece (Russo and Rana 1978; Rana and Kyriakopoulou 1980; Gallitelli et al. 2004; Rana et al. 2005). The virus-infected globe artichoke plants show vein yellowing, yellow flecking, and line patterns on the leaves (Gigante 1951). The virus is transmitted by several aphid species such as Myzus persicae and Aphis fabae in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Broad bean wilt virus Taxonomic position Genus: Fabavirus

(BBWV)

Family: Secoviridae

BBWV (but without specifying either of the currently recognized species BBWV-1 or BBWV-2) infection in plants of Cynara cardunculus var. scolymus was reported from Southern Italy, France, Greece, and Spain (Russo and Rana 1978; Rana et al. 1987b, 2005; Kyriakopoulou 1995; Gallitelli et al. 2004). The virus-infected globe artichoke plants show symptoms of yellow mottle, mosaic, or line patterns. The virus is transmitted by the aphid vector, Capitophorus horni, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BBWV, refer to Vicia faba.

Chinese artichoke mosaic virus Taxonomic position Genus: Potyvirus

(ChAMV)

Family: Potyviridae

Geographical distribution ChAMV infection in plants of Chinese artichoke was reported from Japan (Fuji et al. 2003).

C

794

Cynara cardunculus var. scolymus (Globe artichoke)

Symptoms and host(s) The virus-infected globe artichoke plants exhibit mosaic symptoms. Transmission The virus is transmitted by the aphid vector, Myzus persicae, in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments 680–900 nm in length and 13 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA. A partial genome sequence of 1675 nt is available (AB099711) (Fuji et al. 2003; Revers and Garcia 2015; Wylie et al. 2017).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Cynara cardunculus var. scolymus was reported from Italy, Greece, France, Spain, Tunisia, Turkey, and Slovenia (Migliori et al. 1984b; Chabbouh and Cherif 1990; Baricevich et al. 1995; Paradies et al. 2000; Gallitelli et al. 2004; Ortega et al. 2005; Ergun et al. 2013). The virusinfected globe artichoke plants exhibit symptoms of chlorotic mottling, narrowing of the leaves, leaf deformation, necrosis, and stunting (Migliori et al. 1984b; Paradies et al. 2000). The virus is transmitted by several aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Cynara virus

(CraV)

Taxonomic position CraV is a tentative member of the genus Cytorhabdovirus, in the family Rhabdoviridae. Geographical distribution CraV infection in plants of Cynara cardunculus var. scolymus was reported from Spain and Italy (PenaIglesias et al. 1972; Russo et al. 1975; Rana et al. 1988, 2005; Gallitelli et al. 2004). Symptoms and host(s) CraV has always been found in mixed infections with either a potyvirus or an isometric virus, and therefore no specific symptoms can be attributed to the rhabdovirus. Transmission The virus is transmitted by mechanical sap-inoculation. The experimental host range is essentially restricted to the Solanaceae (Rana et al. 1988). Virion properties and genome The virus particles are bacilliform and enveloped and measure 260  75 nm. The genome is not segmented and contains a single molecule of linear single-stranded RNA. No sequence is currently available.

Cynara cardunculus var. scolymus (Globe artichoke)

Pelargonium zonate spot virus Taxonomic position Genus: Anulavirus

795

(PZSV)

Family: Bromoviridae

PZSV infection in plants of Cynara cardunculus var. scolymus was reported from Italy, Europe, and the Mediterranean Basin (Rana and Martelli 1983; Gallitelli et al. 2004; Rana et al. 2005). The leaves of the virus-infected globe artichoke plants exhibit chlorotic mottling, and the plants are stunted. Thrips (Melanothrips fuscus) are involved in transmitting this virus by carrying the virus-infected pollen grains on their bodies and introducing the virus into the susceptible hosts while feeding. The virus is mechanically sap-transmissible to a number of test plants. For more details of PZSV, refer to Pelargonium spp.

Potato virus X Taxonomic position Genus: Potexvirus

(PVX)

Family: Alphaflexiviridae

PVX infection in plants of Cynara cardunculus var. scolymus was reported from Tunisia, Europe, and the Mediterranean Basin (Chabbouh 1989; Chabbouh et al. 1990; Gallitelli et al. 2004; Rana et al. 2005). The virus-infected globe artichoke plants exhibit symptoms of mosaic, leaf deformation, and a strong decrease in leaf size. No vector is identified for this virus. The virus spreads through contact between plants. The virus is mechanically transmissible to herbaceous hosts. For more details of PVX, refer to Solanum tuberosum.

Raspberry ringspot virus Taxonomic position Genus: Nepovirus

(RpRSV)

Family: Secoviridae

RpRSV infection in plants of Cynara cardunculus var. scolymus was reported from Greece (Rana et al. 1985; Roca et al. 1986). The virus-infected globe artichoke plants exhibit yellow blotches on their leaves. The virus is transmitted by nematode vectors (Longidorus spp.) in a non-persistent manner, and also by mechanical sap-inoculation. Vegetative propagation from virus-infected plants is the primary factor for the spread of this virus. For more details of RpRSV, refer to Rubus spp.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV-infected Cynara cardunculus var. scolymus plants were reported from Italy and Turkey (Gallitelli et al. 2004; Rana et al. 2005). These virus-infected globe artichoke plants are symptomless. The virus is known for its highly infectious property, without any known vector. The virus is mechanically saptransmissible and also through contact between plants. For more details of TMV, refer to Nicotiana tabacum.

C

796

Cynara cardunculus var. scolymus (Globe artichoke)

Tobacco rattle virus

(TRV)

Taxonomic position Genus: Tobravirus

Family: Virgaviridae

TRV infection in plants of Cynara cardunculus var. scolymus was reported from Brazil, Europe, and the Mediterranean Basin (Chagas and Silberschmidt 1972; Chagas et al. 1969; Migliori and Marzin 1985; Migliori et al. 1985; Gallitelli et al. 2004; Rana et al. 2005). The virus-infected globe artichoke plants exhibit bright yellow discoloration of the leaves. The virus is transmitted by a nematode vector, Trichodorus christiei. The virus is seed-transmitted in artichoke (Salomao 1976). The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV was reported in plants of Cynara cardunculus var. scolymus in Brazil, Europe, and the Mediterranean Basin (Costa and Tasaka 1971; Gallitelli et al. 2004; Rana et al. 2005). The virus-infected globe artichoke plants exhibit symptoms of stunting and malformed leaves. The virus is transmitted in nature by thrips (Thrips tabaci, Frankliniella occidentalis). The thrips vectors assist in the transfer of virusinfected pollen to the healthy plants. The virus is mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato black ring virus Taxonomic position Genus: Nepovirus

(TBRV)

Family: Secoviridae

TBRV infection in plants of Cynara cardunculus var. scolymus was reported from Europe and the Mediterranean Basin (Migliori et al. 1984a; Rana et al. 1985, 1987a; Murant et al. 1996; Gallitelli et al. 2004). The virus-infected globe artichoke plants exhibit systemic chlorotic leaf spots on the leaves. In some artichoke cultivars, mild mottling of leaves was noticed (Migliori et al. 1987). The total number of plant heads and weight was reduced. The virus is transmitted by the nematode vector, Longidorus attenuatus, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TBRV, refer to Solanum lycopersicum.

Tomato infectious chlorosis virus Taxonomic position Genus: Crinivirus

(TICV)

Family: Closteroviridae

TICV-infected plants of Cynara cardunculus var. scolymus were reported from Italy and California (USA) (Gallitelli et al. 2004; Davino et al. 2009; Salleh et al. 2014). In virus-infected globe artichoke

Cynara cardunculus var. scolymus (Globe artichoke)

797

plants there may be either no clear symptoms or interveinal yellowing of the leaves, depending on the cultivar. The virus is transmitted by the whitefly vector, Trialeurodes vaporariorum, in a semipersistent manner (Duffus et al. 1996). The virus is not transmissible by mechanical sap-inoculation. For more details of TICV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Cynara cardunculus var. scolymus was reported from Argentina, Australia, Greece, Turkey, Italy, Spain and California (USA) (Gracia and Feldman 1978; Martelli et al. 1981; Gallitelli et al. 2004; Camele and Rana 2005; Ortega et al. 2005; Vilchez et al. 2005; Lotos et al. 2009; Testa et al. 2011). The virus-infected globe artichoke plants exhibit severe necrosis with leaf curl, stunted growth, generalized chlorosis, and necrotic spots extending to leaf blades, leaf and capitulum distortion, and general wilting of the plant that often results in death, especially if the plant becomes infected in the early growth stages. Infected plants exhibit gradual vascular bundle browning visible in a stem cross section. The virus is transmitted by thrips vectors, Frankliniella occidentalis and Thrips tabaci, in a persistent-propagative manner (Gallitelli 2004) and also by mechanical sap-inoculation to a large number of vegetable and ornamental crops. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Cynara cardunculus var. scolymus was reported from Italy (Foddai et al. 1993; Gallitelli et al. 2004; Rana et al. 2005). This virus-infected globe artichoke plants are symptomless. The virus is transmitted by aphid vectors such as Brevicoryne brassicae and Myzus persicae in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

References Baricevich D, Kus M, Pepelnjak M (1995) Isolation of IgG immunoglobulins from rabbit ALV-antiserum using proteinA-sefarose and estimation of globe artichoke local cultivar “Istra” for the presence of Artichoke latent virus (ALV) and Cucumber mosaic virus (CMV) in Slovenia. Zbornik Biotechniske Fakultete Univerze Ljubijani, Kmetijstvo 65:47–54 Bottalico G, Padula M, Campanale A, Finetti-Sialer MM, Saccomanno F, Gallitelli D (2002) Seed transmission of Artichoke Italian latent virus and Artichoke latent virus in globe artichoke. J Plant Pathol 84:167–168 Brown DJF, Trudgill DL (1998) Nematode transmission of plant viruses: a 30 year perspective. Host pathogen interactions and crop protection. SCRI Annual Report. pp 121–125

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Cynara cardunculus var. scolymus (Globe artichoke)

Brown DJF, Kyriakopoulou PE, Robertson WM (1997) Frequency of transmission of Artichoke Italian latent nepovirus by Longidorus fasciatus (Nematoda: Longidoridae) from artichoke fields in the Iria and Kandia areas of Argopolis in northeast Peloponnesus, Greece. Eur J Plant Pathol 103:501–506 Camele I, Rana GL (2005) New outbreaks of Tomato spotted wilt tospovirus (TSWV) infections on globe artichoke in Basilicapa and Apulia. Acta Hortic 681:593–595 Chabbouh N (1989) Mise en evidence de quatre virus presents sur l’artichaut en Tunisie. Ann I’Istitut Natl Rech Agronomique Tunis 62:3–14 Chabbouh N, Cherif C (1990) Outbreaks and new records. Tunisia. Cucumber mosaic virus in artichoke. FAO Plant Protect Bull 35:52–53 Chabbouh N, Cherif C, Martelli GP (1990) Natural infections of artichoke by Potato virus X in Tunisia. J Phytopathol 129:257–260 Chagas CM, Silberschmidt KM (1972) Virus da faixa amarela da alcachofra: occorencia, transmissao mecanica e propriedades fisicas. Biologico 38:35–40 Chagas CM, Flores M, Carner J (1969) Una nova doenca de virus da alcahofra no Estado de Sao Paulo. Biologico 35:271–274 Ciuffo M, Testa M, Lenzi R, Turina M (2011) Ranunculus latent virus: A strain of artichoke latent virus or a new macluravirus infecting artichoke? Arch Virol 156:1053–1057 Costa AS, Tasaka H (1971) Enfezamento e malformacao foliar de alcachofra inducido pelo virus da necrose branca do fumo. Biologico 37:176–179 Costa AS, Duffus JE, Morton DJ, Iarwood CE, Bardin R (1959) A latent virus of California artichokes. Phytopathology 49:49–53 Davino S, Tomassoli L, Tiberini A, Mondello V, Davino M (2009) Outbreak of Tomato infectious chlorosis virus in a relevant artichoke producing area of Sicily. J Plant Pathol 91(Suppl. 4):S4.57–S4.58 Di Franco A, Gallitelli D, Vovlas C, Martelli GP (1989) Partial characterization of Artichoke virus M. J Phytopathol 127:265–273 Duffus JE, Liu H-Y, Wisler GC (1996) Tomato infectious chlorosis virus – a new clostero-like virus transmitted by Trialeurodes vaporariorum. Eur J Plant Pathol 102:219–226 Dunez J, Le Gall O (2011) Nepovirus. Comoviridae. In: The Springer index of Viruses. Springer, New York, pp 361–369. https://doi.org/10.1007/978-0-387-95919-1_55 Elbeaino T, Belghacem I, Mascia T, Gallitelli D, Digiaro M (2017) Next generation sequencing and molecular analysis of artichoke Italian latent virus. Arch Virol 162:1805–1809 Ergun M, Erkan S, Paylan IC (2013) Cucumber mosaic virus in globe artichoke in Turkey. Can J Plant Pathol 35:514–517 Erkan S, Gumus M, Duman I, Can Paylan I, Ergun M (2014) The new report of artichoke latent virus (ArLV) from globe artichoke in Turkey. Ege Univ Ziraat Fak Derg 51:265–269 Fischer HV, Lockhart BE (1974) A Moroccan isolate of Artichoke latent virus. Plant Dis Rptr 58:1123–1126 Foddai A, Marras F, Idini G (1977) Presenza di un “Potyvirus” sul Carciofo (Cynara scolymus L.) in Sardegna. Ann Facolta Agrar Univ Sassari 25:398–407 Foddai A, Corda P, Idini G (1983) Influenza del potyvirus latente del carciofo spinoso sardo sulla produttività delle piante in pieno campo. II. Risultati relativi al secondo anno di impianto. Ann Facoltà Agraria Univ Sassari 30:37–43 Foddai A, Cugusi M, Idini G (1993) Artichoke (Cynara scolymus L.): a new host for Turnip mosaic virus. Phytopathol Mediterr 32:247–248 Foster GD (2011) Macluravirus. Potyviridae. In: The Springer index of Viruses. Springer, New York, pp 1421–1424. https://doi.org/10.1007/978-0-387-95919-1_234 Fuji S, Yamamoto H, Furuya H, Naito H (2003) Characterization of a new potyvirus isolated from Chinese artichoke in Japan. Arch Virol 148(11):2249–2255 Gallitelli D (2004) The most important virus diseases of vegetable crops in open field. Informatore Fitopatologico 54:25–29 Gallitelli D, Rana GL, Vovlas C, Martelli GP (2004) Viruses of globe artichoke: an overview. J Plant Pathol 86:267–281 Gallitelli D, Mascia T, Martelli GP (2012) Viruses in artichoke. Adv Virus Res 84:289–324 Gigante R (1951) Il mosaico del carciofo. Boll Staz Pat Veg Roma 7:177–181 Gracia O, Feldman JL (1978) Natural infection of artichoke by Tomato spotted wilt virus. Plant Dis Rptr 62:1076–1077 King AMQ, Adams MJ, Carstens EB, Lefkowitz EJ (2012) Virus taxonomy: classification and nomenclature of viruses, Ninth Report of the International Committee on Taxonomy of Viruses. Elsevier Academic Press, San Diego Kyriakopoulou PE (1995) Artichoke Italian latent virus causes artichoke patchy chlorotic stunting disease. Ann Appl Biol 127:489–497 Kyriakopoulou PE, Bem FP (1973) Some virus diseases of cultivated plants noticed in Greece in the years 1971 and 1972. First symposium of geotechnical research B.I. pp 409–418 Kyriakopoulou PE, Bem FP (1982) Artichoke Marathon yellow blotch, a new virus disease of artichoke. Benaki Phytopathological Institute, Annual Report 1982. pp 90–91

Cynara cardunculus var. scolymus (Globe artichoke)

799

Kyriakopoulou PE, Rana GL, Roca F (1985) Geographical distribution, natural host range, pollen and seed transmissibility of Artichoke yellow ring spot virus. Ann I’Inst Phytopathologique Benaki, NS 14:139–155 Lopez-Moya JJ, Valli A, Garcia JA (2009) Potyviridae. In: Encyclopaedia of life sciences (ELS). Iacobellis NS, Collmer A, Hutcheson SW, Mansfield JW, Morris CE, Murillo J, Schaad D NW, Stead E, Surico G, Ullrich M (eds). Kluwer Academic, Dordrecht, The Netherlands. https://doi.org/10.1002/9780470015902.a0000755.pub2 Lotos L, Efthimiou K, Chatzivassiliou EK, Dizmou D, Katis NI (2009) First report of Tomato spotted wilt virus in globe artichoke in Greece. J Plant Pathol 91:97–112 Majorana G, Rana GL (1970a) Un nuovo virus latente isolate da carciofo in Puglia. Phytopathol Mediterr 9:193–196 Majorana G, Rana GL (1970b) A latent virus of artichoke belonging to Potato virus S group. Phytopathol Mediterr 9:200–202 Marrou J, Mehani S (1964) Etude d’un virus parasite de I’artichaut. C Rendues Acad Agric Fr 50:1051–1064 Martelli GP, Rana GL (1976) Viruses and virus diseases of globe artichoke and cardoon. Nuovi studi sul Carciofo. Proceedings of the 2nd Congresso Internazionale sul Carciofo, Bari 1973. pp 811–830 Martelli GP, Russo M, Rana GL (1976) Occurrence of Artichoke mottled crinkle virus in Malta. Plant Dis Reptr 60:13–133 Martelli GP, Russo M, Rana GL (1981) A survey of the virological problems of Cynara species. Studi sul Carciofo Atti del  3 Congresso Internazionale sul Carciofo, Bari. pp 895–927 Migliori A, Marzin H (1985) Presence du Tobacco rattle virus (TRV) dans les cultures d’artichaut en France. Agronomie 5:549–552 Migliori A, Marzin H, Rana GL (1984a) Mise en evidence du tomato black ring virus (TBRV) chez l’artichaut en France. Agronomie 4:683–686 Migliori A, Lot H, Pecaut P, Duteil M, Rouze-Jouan J (1984b) Les virus de I’artichaut. I. mise en evidence de trois virus dans les cultures francoises d’artichaut. Agronomie 4:257–268 Migliori A, Marzin H, Legal V, Homo E, Corre J (1985) Presence du tobacco rattle virus (TRV) dans les cultures d’artichaut en France. Agronomie 5:549–552 Migliori A, Homo E, Corre J, Marzin H, Legal V, Curvale JP (1987) Repartition, frequence et nuisibilite des virus chez I’artichaut en Bretagne. PHM Rev Hortic 247:29–36 Minutillo SA, Marais A, Mascia T, Faure C, Svanella-Dumas L, Thail S, Payet A, Perennec S, Schoen L, Gallitelli D, Candresse T (2015) Complete nucleotide sequence of artichoke latent virus shows it to be a member of genus Macluravirus in the family Potyviridae. Phytopathology 105:1155–1160 Murant AF, Jones AT, Martelli GP, Stace-Smith R (1996) Nepoviruses: general properties, diseases and virus identification. In: Harrison BD, Murant AF (eds) The plant viruses: polyhedral virions and bipartite RNA genomes. Plenum Press, New York, pp 99–137 Ortega AM, Juarez M, Jorda MC, Armengol J (2005) Viral diseases in artichoke crops in Spain. Acta Hortic 681:611–616 Paradies F, Finetti Sialer M, Di Franco A, Gallitelli D (2000) First report on the occurrence of Cucumber mosaic virus in artichoke in Italy. J Plant Pathol 82:244 Paylan IC, Ergun M, Erkan S (2013) First report of Artichoke yellow ringspot virus in globe artichoke in Turkey. Plant Dis 97:1388 Pena-Iglesias A, Rubio-Huertos M, Moreno San Martin R (1972) Un virus de tipo bacilliforme en alcachofra (Cynara scolymus). Ann Inst Nac Investig Agron 2:123–137 Rana GL, Cherif C (1981) Occurrence of Artichoke mottled crinkle virus in Tunisia. Phytopathol Mediterr 20:179–180 Rana GL, Kyriakopoulou PE (1980) Bean yellow mosaic virus in artichokes in Greece. Proceedings 5th Congress of the Mediterranean Phytopathological Union, Patras 1980. pp 38–40 Rana GL, Kyriakopoulou PE (1982) Artichoke Italian latent and artichoke mottled crinkle viruses in artichoke in Greece. Phytopathol Medit 22:46–48 Rana GL, Martelli GP (1983) Virosi del carciofo. Italia Agric 120:27–38 Rana GL, Gallitelli D, Kyriakopoulou PE, Russo M, Martelli GP (1980) Host range and properties of Artichoke yellow ring spot virus. Ann Appl Biol 97:177–185 Rana GL, Russo M, Gallitelli D, Martelli GP (1982) Artichoke latent virus: characterization, ultrastructure, and geographical distribution. Ann Appl Biol 101:179–188 Rana GL, Castrovilli S, Gallitelli D, Kyriakopoulou PE (1985) Studies on two serologically distinct raspberry ringspot virus strains from artichoke. Phytopathol Z 112:222–228 Rana GL, Di Franco A, Piazzolla P, Migliori A (1987a) Further studies on a Tomato black ring virus isolate from artichoke. J Phytopathol 118:203–211 Rana GL, Migliori A, Ragozzino A (1987b) La maculatura gialla del carciofo in Campania e Puglia. Inf Fitopatol 37:41–43 Rana GL, Di Franco A, Galasso I (1988) Further studies on Cynara rhabdovirus. J Phytopathol 123:147–155 Rana GL, Piazzolla P, Lafortezza R, Greco N (1989) Characterization of a latent elongated virus from globe artichoke (Cynara scolymus L.) in Italy. J Phytopathol 125:289–298

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Cynodon dactylon (Bermuda grass)

Rana GL, Elia A, Nuzzaci M, Lafortezza R (1992) Effect of artichoke latent virus infection on the production of artichoke heads. J Phytopathol 135:153–159 Rana GL, Gallitelli D, Vovlas C, Martelli GP (2005) Viruses of globe artichoke: an overview. Acta Hortic 681:555–572 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Roca F, Martelli GP, Lamberti F, Rana GI (1975) Distribution of Longidorus attenuatus Hooper in Apulian artichoke fields and its relationship with Artichoke Italian latent virus. Nematol Mediterr 3:91–101 Roca F, Rana GL, Kyriakopoulou PE (1982) Longidorous fasciatus Roca et Lamberti vector of a serologically distinct strain of Artichoke Italian latent virus in Greece. Nematol Mediterr 10:65–69 Roca F, Rana GL, Kyriakopoulou PE (1986) Studies on Longidoridae (Nematoda: Dorylaimida) and Raspberry ringspot virus spread in some artichoke fields in Greece. Nematol Mediterr 14:251–256 Russo M, Rana GL (1978) Occurrence of two legume viruses in artichoke. Phytopathol Mediterr 17:212–216 Russo M, Martelli GP, Rana GL (1975) A rhabdovirus of Cynara in Italy. Phytopathol Z 83:223–231 Salleh W, Mnari-Hattab M, Minutillo SA, Spano R, Zammouri S, Gallitelli D (2014) First report of Tomato infectious chlorosis virus in Tunisia. J Plant Pathol 96:433 Salomao T (1976) Soil transmission of Artichoke yellow band virus. Studi sul Carciofo Atti del 2 Congresso Internazionale sul Carciofo, Bari 1973. pp 831–854 Sanfacon H (2015) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. In: Encyclopedia of Life Sciences. Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000764.pub3. http:// www.els.net Sanfacon H, Wellink J, Le Gall O, Karasev A, van der Vlugt R, Wetzel T (2009) Secoviridae: a proposed family of plant viruses within the order Picornavirales that combines the families Sequiviridae and Comoviridae, the unassigned genera Cheravirus and Sadwavirus, and the proposed genus Torradovirus. Arch Virol 154(5):899–907 Savino V, Gallitelli D, Jankulova M, Rana GL (1977) A comparison of four isolates of Artichoke Italian latent virus (AILV). Phytopathol Mediterr 16:41–50 Tavazza M, Lucioli A, Calogero A, Pay A, Tavazza R (1994) Nucleotide sequence, genomic organization and synthesis of infectious transcripts from a full-length clone of artichoke mottle crinkle virus. J Gen Virol 75:1515–1524 Testa M, Sanna D, Pintore R, Marongiu G, Marras PM (2011) A two-year survey of TSWVoccurrence in globe artichokes in Sardinia (ITALY). Acta Hortic (ISHS) 917:297–302 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J GenVirol 98:529–531 Vilchez M, Paulus AO, Moyer JW, Sehrader WL (2005) First report of Tomato spotted wilt virus affecting globe artichoke in California, USA. Acta Hortic 681:607–610 White KA (2011) Tombusvirus. Tombusviridae. In: The Springer index of Viruses. Springer, New York, pp 1917–1926. https://doi.org/10.1007/978-0-387-95919-1_314 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Cynodon dactylon (Bermuda grass) Family: Poaceae

Forage crop

Barley yellow dwarf virus MAV Genus: Luteovirus

(Acronym: BYDV-MAV)

Family: Luteoviridae

BYDV-MAV infection in plants of Cynodon dactylon was reported from Western Australia (Jones et al. 1990; Hawkes and Jones 2005). The virus is transmitted by number of aphid species in a circulative, non-propagative manner. No mechanical transmission of this virus is reported. For more details of BYDV-MAV, refer to Hordeum vulgare.

Cynodon dactylon (Bermuda grass)

Barley yellow dwarf virus PAV Taxonomic position Genus: Luteovirus

801

(BYDV-PAV)

Family: Luteoviridae

BYDV-PAV infection in plants of Cynodon dactylon was reported from Western Australia and Syria (Jones et al. 1990; Hawkes and Jones 2005; Ansi et al. 2007). The virus is transmitted by a number of aphid species in a circulative, non-propagative manner. No mechanical transmission of this virus is reported. For more details of BYDV-PAV, refer to Hordeum vulgare.

Bermuda grass etched-line virus Taxonomic position Genus: Marafivirus

(BELV)

Family: Tymoviridae

Geographical distribution BELV infection was first reported in plants of Cynodon dactylon and Sorghum halepense from Morocco by Lockhart et al. (1985a). The virus spreads in Morocco and Iran (Izadpanah 1986; Izadpanah et al. 2000). Symptoms and host(s) The virus-infected Bermuda grass plants exhibit symptoms of white etched lines and spots in leaves and are stunted. The natural hosts of this virus are Cynodon dactylon, Echinochloa colonum, and Sorghum halepense. Transmission The virus is transmitted by a leafhopper vector, Aconurella prolixa, in a persistent propagative manner. The virus is not transmissible by mechanical inoculation. The virus is not transmitted by seed (Lockhart et al. 1985a). Virion properties and genome The virions are non-enveloped, isometric capsid with T = 3 icosahedral symmetry, and about 30 nm in diameter. The genome is a monopartite, linear, positive-sense, single-stranded RNA of approx 6000–7000 nts. There is no VPg at the 50 -terminus. A partial genome sequence of 1093 nt is available (AY040531) (Masumi and Izadpanah 1991, 1996; Izadpanah et al. 2000; Martelli et al. 2002; Hammond et al. 2011).

Bermuda grass mosaic virus

(BGMV)

Taxonomic position BGMV is a tentative member of the genus Potyvirus and family Potyviridae Geographical distribution BGMV infection in plants of Cynodon dactylon was reported from Iran (Hosseini et al. 2010).

C

802

Cynodon dactylon (Bermuda grass)

Symptoms and host range The virus-infected plants exhibit mosaic symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner, and the virus is also mechanically sap-transmissible. Virion properties and genome The virions are flexuous filaments. The genome is positive-sense single-stranded RNA. A partial genome sequence of 1668 nt is available (GU552269) (Wylie et al. 2017).

Bermuda grass southern mosaic virus

(BGSMV)

Taxonomic position BGSMV is a tentative member of the genus Potyvirus and family Potyviridae Geographical distribution BGSMV infection in plants of Cynodon dactylon was reported from Iran (Bragard and Steyer 2005). Symptoms and host(s) The virus-infected Bermuda grass plants exhibit mosaic symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. Virion properties and genome The virions are flexuous filaments. The genome is a single molecule of positive-sense single-stranded RNA of 9589 nt (KU372146) (Wylie et al. 2017).

Cereal yellow dwarf virus RPV

(CYDV-RPV)

Synonyms Barley yellow dwarf virus RPV Taxonomic position Family: Luteoviridae

Genus: Polerovirus

CYDV-RPV infection in plants of Cynodon dactylon was reported from Western Australia (Jones et al. 1990; Hawkes and Jones 2005). The virus is transmitted by number of aphid species in a circulative, non-propagative manner. No mechanical transmission of this virus is reported. For more details of CYDV-RPV, refer to Hordeum vulgare.

Cynodon chlorotic streak virus

(CCSV)

Taxonomic position CCSV is a tentative member of the genus Nucleorhabdovirus and family Rhabdoviridae

Cynodon dactylon (Bermuda grass)

803

Geographical distribution CCSV infection in plants of Cynodon dactylon was reported from France, Iran, Jordan, Morocco, Spain, and Tunisia (Lockhart et al. 1985b; Lamprecht et al. 2009). Symptoms and host(s) The virus-infected Bermuda grass plants exhibit chlorotic streaking symptoms. Transmission The virus is transmitted by the planthopper vector, Toya propinqua, in a persistent, propagative manner. The virus is not transmissible by mechanical inoculation nor is it transmitted by contact between plants. Virion properties and genome The virions are rhabdo- or bullet-shaped, enveloped, usually straight, with a clear modal length of 240 nm (in sap) or 280 nm (in thin sections), and 72 nm wide (in sap) or 80 nm wide (in thin sections). The genome is presumed to be a single negative stranded RNA of >12 kb.

Cynodon rhabdovirus

(CRV)

Taxonomic position CRV is a tentative member of the genus Nucleorhabdovirus and family Rhabdoviridae Geographical distribution CRV infection in plants of Cynodon dactylon was reported from South Africa (Lamprecht et al. 2009). Virion properties and genome The virions are bullet shaped and measures 63 nm in diameter and 240 nm in length. A partial genome sequence is available (EU650683) (Walker et al. 2018).

Grapevine fanleaf virus Taxonomic position Genus: Nepovirus

(GFLV)

Family: Secoviridae

GFLV infection in plants of Cynodon dactylon was reported from Iran (Izadpanah et al. 2003). The virus-infected Bermuda grass plants exhibit symptoms of systemic vein-clearing, chlorotic spotting, and leaf distortion. The virus is transmitted by a nematode vector Xiphinema index in a nonpersistent manner, and also by mechanical sap-inoculation. For more details of GFLV, refer to Vitis vinifera.

Maize yellow dwarf virus RMV Synonyms Barley yellow dwarf virus RMV

(MYDV-RMV)

C

804

Cynodon dactylon (Bermuda grass)

Taxonomic position Genus: Polerovirus

Family: Luteoviridae

MYDV-RMV infection in plants of Cynodon dactylon was reported from Western Australia (Jones et al. 1990; Hawkes and Jones 2005). The virus is transmitted by number of aphid species in a circulative, non-propagative manner. No mechanical transmission of this virus is reported. For more details of MYDV-RMV, refer to Hordeum vulgare.

Sugarcane mosaic virus Taxonomic position Genus: Potyvirus

(SCMV)

Family: Potyviridae

SCMV infection in plants of Cynodon dactylon and C. nlemfuensis was reported from Kenya (Louie 1980). The virus-infected Bermuda grass plants exhibit mosaic symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SCMV, refer to Saccharum officinarum.

References Ansi A, Kumari SG, Haj Kasem A, Makkouk KM, Muharram I (2007) The Occurrence of Barley yellow dwarf viruses on cereal crops and Wild Grasses in Syria. Arab J Pl Prot 25:1–9 Bragard C, Steyer S (2005) Bermuda grass southern mosaic virus: a distinct Potyvirus infecting several gramineous species in Iran. Parasitica 61:105–110 Hammond RW, Edwards MC, Ramirez P (2011) Marafivirus. Tymoviridae. In: The Springer index of Viruses. Springer, New York, pp 1947–1952. https://doi.org/10.1007/978-0-387-95919-1_319 Hawkes JR, Jones RAC (2005) Incidence and distribution of Barley yellow dwarf virus and Cereal yellow dwarf virus in over-summering grasses in a Mediterranean-type environment. Aust J Ag Res 56:257–270 Hosseini A, KoohiHabibi M, Izadpanah K, Mosahebi GH, Rubies-Autonell C, Ratti C (2010) Characterization of a filamentous virus from Bermuda grass and its molecular, serological and biological comparison with Spartina mottle virus. Arch Virol 155(10):1675–1680 Izadpanah K (1986) Report of two Bermuda grass viruses from Iran. Proc 9th Plant Protec Cong Iran Ferdowsi Univ, Mashhad, Iran. p 161 Izadpanah K, Zhang YP, Masumi M, Rowhani A (2000) Cloning and sequencing of the Iranian isolate of Bermuda grass etched-line virus. Phytopathology 90:538 Izadpanah K, Zaki-Aghl M, Zhang YP, Daubert SD, Rowhani A (2003) Bermuda grass as a potential reservoir host for Grapevine fanleaf virus. Plant Dis 87:1179–1182 Jones RAC, McKirdy SJ, Shivas RG (1990) Occurrence of barley yellow dwarf viruses in over-summering grasses and cereal crops in Western Australia. Australas Plant Pathol 19:90–96 Lamprecht RL, Pietersen G, Kasdorf GGF, Nel LH (2009) Characterisation of a proposed Nucleorhabdovirus new to South Africa. Eur J Plant Pathol 123:105–110 Lockhart BEL, Khaless J, Lennon AM, EL Maatoui M (1985a) Properties of Bermuda grass etched-line virus, a new leafhopper transmitted virus related to Maize rayado fino and Oat blue dwarf viruses. Phytopathology 75:1258–1262 Lockhart BEL, Khaless N, El Maataoui M, Lastra R (1985b) Cynodon chlorotic streak virus, a previously undescribed plant rhabdovirus infecting Bermuda grass and maize in the Mediterranean area. Phytopathology 75:1094–1098 Louie R (1980) Sugarcane mosaic virus in Kenya. Plant Dis 64:944–947 Martelli GP, Sabanadzovic S, Abou-Ghanem Sabanadzovic N, Edwards MC, Dreher T (2002) The family Tymoviridae. Arch Virol 147(9):1837–1846 Masumi M, Izadpanah K (1991) Purification and physicochemical properties of Bermuda grass etched-line virus (Iranian isolate). Iran J Plant Pathol 27:96–97 Masumi M, Izadpanah K (1996) Properties of the Iranian isolate of Bermuda grass etched-line virus. J Phytopathol 144:231–234

Cynosurus cristatus (Crested dog’s-tail)

805

Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Rhabdoviridae. J GenVirol 99:447–448 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

C Cynoglossum officinale (Houndstongue) Family: Boraginaceae

Medicinal

Alfalfa mosaic virus

(AMV)

Taxonomic position Genus: Alfamovirus

Family: Bromoviridae

AMV infection in plants of Cynoglossum officinale was reported from Italy (Bellardi et al. 2002). The virus-infected houndstongue plants show stunting, interveinal chlorotic spots, and midrib necrosis symptoms. The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

References Bellardi MG, Rubies-Autonell C, Pisi A (2002) Cynoglossum officinale, a new natural host of Alfalfa mosaic virus. Plant Dis 86:444

Cynosurus cristatus (Crested dog’s-tail) Family: Poaceae

Forage crop

Cynosurus mottle virus

(CnMoV)

Taxonomic position CnMoV is a tentative member of the genus Sobemovirus and family Solemoviridae Geographical distribution CnMoV infection in plants of Cynosurus cristatus was reported from Germany, New Zealand, and the UK (Mohamed 1978; Mohamed and Mossop 1981). Symptoms and host(s) The virus-infected Crested dog’s-tail plants exhibit severe leaf mottling and necrosis symptoms. The natural host range of this virus are Agrostis tenuis, A. stolonifera, Cynosurus cristatus, and Lolium perenne (Catherall et al. 1977).

806

Cyperus spp. (Nutsedge)

Transmission The virus is reported to be transmitted by the aphid vector Rhopalosiphum padi in New Zealand (Mohamed 1978) and by the cereal leaf beetle Oulema melanopus in the UK. The virus is mechanically sap-transmissible. Virion properties and genome The virions are isometric with a diameter of 28 nm. The genome is a positive-sense single-stranded RNA (Hull 1988; Somera et al. 2015). No sequence is currently available.

Wheat dwarf virus Taxonomic position Genus: Mastrevirus

(WDV)

Family: Geminiviridae

WDV infection in plants of Cynosurus cristatus was reported from the Czech Republic (Ripl and Kundu 2015). The virus is transmitted by the leafhopper vector, Psammotettix alienus, in a persistent, circulative, and non-propagative manner. The virus is not transmissible by mechanical inoculation, and not transmitted by contact between plants. The virus is not transmitted by seed and pollen. For more details of WDV, refer to Triticum aestivum.

References Catherall PL, Andrews EA, Chamberlain JA (1977) Host-ranges of cocksfoot mottle and cynosurus mottle viruses. Ann Appl Biol 87:233–235 Hull T (1988). The sobemovirus group. In: Koenig R(ed) The plant viruses. Polyhedral virions with monopartite RNA genomes, vol 3. Plenum Press, New York, pp 113–146. Mohamed NA (1978) Cynosurus mottle virus, a virus affecting grasses in New Zealand. N Z J Agric Res 21:709–714 Mohamed NA, Mossop DW (1981) Cynosurus and cocksfoot mottle viruses: a comparison. J Gen Virol 55:63 Ripl J, Kundu JK (2015) Cynosurus cristatus, a new host of Wheat dwarf virus in the Czech Republic. J Plant Pathol 97:547 Somera M, Sarmiento C, Truve E (2015) Overview on Sobemoviruses and a proposal for the creation of the family Sobemoviridae. Viruses 7:3076–3115

Cyperus spp. (Nutsedge) Family: Cyperaceae

Weed host

Brome streak mosaic virus Taxonomic position Genus: Tritimovirus

(BrSMV)

Family: Potyviridae

BrSMV infection in plants of Cyperus esculentus was reported from Hungary (Takacs et al. 2008). The virus is transmitted by the mite vector in a semi-persistent manner, and also by mechanical sapinoculation. For more details of BrSMV, refer to Bromus spp.

Cyphomandra betacea (Tamarillo, Tree tomato)

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

807

(INSV)

Family: Tospoviridae

INSV infection in plants of Cyperus spp. was reported from Georgia (Martinez-Ochoa et al. 2004). The virus-infected nutsedges plants do not exhibit any symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large a number of hosts. For more details of INSV, refer to Impatiens spp.

References Martinez-Ochoa N, Mullis SW, Csinos AS, Webster TM (2004) First report of yellow nutsedge (Cyperus esculentus) and purple nutsedge (C. rotundus) in Georgia naturally infected with Impatiens necrotic spot virus. Plant Dis 88:771 Takacs AP, Kazinczi G, Horvath J, Gaborjanyi R (2008) Cyperuses culentus L. a new host of Brome streak mosaic virus (BrSMV). J Plant Dis Prot 21:527–528

Cyphomandra betacea (Tamarillo, Tree tomato) Synonyms Solanum betaceum Family: Solanaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Edible fruit

(AMV)

Family: Bromoviridae

AMV infection in plants of Solanum betaceum was reported from Colombia and New Zealand (Fletcher 1987; Eagles et al. 1994; Jaramillo et al. 2011). The virus-infected tamarillo plants exhibit bright yellow mosaic and vein-banding symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

ArMV infection in plants of Cyphomandra betaceum was reported from New Zealand (Thomas and Procter 1972). The virus-infected tamarillo plants exhibit symptoms of sunken necrotic rings on the

C

808

Cyphomandra betacea (Tamarillo, Tree tomato)

fruits. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Cucumber mosaic virus

(CMV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV infection in of Solanum betaceum was reported from Colombia and New Zealand (Eagles et al. 1994; Jaramillo et al. 2011). The virus-infected tamarillo plants exhibit leaf narrowing symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Peru tomato mosaic virus Taxonomic position Genus: Potyvirus

(PTV)

Family: Potyviridae

PTV infection in plants of Solanum betaceum was reported from Ecuador (Insuati et al. 2016). The virus-infected tamarillo plants exhibit mosaic, mottling, and leaf deformation symptoms. The virus is transmitted by aphid vectors in a non-persistent manner only in the presence of a co-infecting potyvirus (Kassanis and Govier 1971) and also by mechanical sap-inoculation, and by grafting. For more details of PTV, refer to Solanum lycopersicum.

Potato aucuba mosaic virus Taxonomic position Genus: Potexvirus

(PAMV)

Family: Alphaflexiviridae

PAMV infection in plants of Cyphomandra betacea was reported from New Zealand (Mossop 1982; Eagles et al. 1994). The virus-infected tamarillo plants do not exhibit any symptoms. The virus is transmitted by the aphid vector, Myzus persicae, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PAMV, refer to Solanum tuberosum.

Potato leafroll virus Taxonomic position Genus: Polerovirus

(PLRV)

Family: Luteoviridae

PLRV infection in plants of Solanum betaceum was reported from Ecuador (Jaramillo et al. 2011; Sivaprasad et al. 2016a). The virus-infected tamarillo plants exhibit symptoms of mosaic, blistering, vein-banding, necrotic rings, leaf deformation, and etching in fruits. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical inoculation. For more details of PLRV, refer to Solanum tuberosum.

Cyphomandra betacea (Tamarillo, Tree tomato)

Potato virus A

809

(PVA)

Synonyms Tamarillo mosaic virus (TamMV) Taxonomic position Genus: Potyvirus

C

Family: Potyviridae

PVA (as TamMV) infection was first reported in plants of Cyphomandra betacea from New Zealand by Chamberlain (1954). The virus spreads in New Zealand (Mossop 1977; Eagles et al. 1990, 1994; Eagles 1994). The virus-infected tamarillo plants exhibit symptoms of vein-banding and mosaic, and fruitblotching. The virus is transmitted by the aphid vector Myzus persicae in a non-persistent manner, and also by mechanical sap sap-inoculation (Mossop 1977; Eagles et al. 1990). For more details of PVA, refer to Solanum tuberosum.

Potato virus V

(PVV)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

PVV infection in plants of Solanum betaceum was reported from Ecuador (Insuati et al. 2016). The virus-infected tamarillo plants exhibit mosaic, mottling, and leaf deformation symptoms. The virus is transmitted by several aphid species in a non-persistent manner, and also by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants. For more details of PVV, refer to Solanum tuberosum.

Potato virus Y

(PVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

PVY infection in plants of Solanum betaceum was reported from Ecuador (Bhargava and Joshi 1959; Sivaprasad et al. 2015; Insuati et al. 2016). The virus-infected tamarillo plants exhibit mosaic, mottling, and leaf deformation symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, by mechanical sap-inoculation to a large number of hosts and by grafting. For more details of PVY, refer to Solanum tuberosum.

Tamarillo leaf malformation virus Taxonomic position Genus: Potyvirus

(TLMV)

Family: Potyviridae

Geographical distribution TLMV infection in plants of Solanum betaceum was reported from Colombia (Garcia et al. 2011; Gutierrez et al. 2015; Villegas et al. 2017).

810

Cyphomandra betacea (Tamarillo, Tree tomato)

Symptoms and host(s) The virus-infected tamarillo plants exhibit leaf malformation symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically saptransmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9720 nt (excluding the 30 poly(A) tract) (KM523548 = NC_026615). The genome contains one large ORF that codes for a polyprotein of 3073 residues and is flanked by 50 and 30 untranslated regions (UTR) of 156 and 315 nt, respectively (Gutierrez et al. 2015; Wylie et al. 2017).

Tomato mosaic virus Taxonomic position Genus: Tobamovirus

(ToMV)

Family: Virgaviridae

ToMV infection in plants of Solanum betaceum was reported from Colombia (Jaramillo et al. 2011). The virus-infected tamarillo plants exhibit symptoms of mosaic, blistering, vein-banding, necrotic rings, leaf deformation, and etching in fruits. No vector is involved in the spread of this virus. This virus is transmissible by mechanical sap-inoculation, and transmissible by grafting and also by contact between plants. For more details of ToMV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Solanum betaceum was reported from Colombia (Jaramillo et al. 2011). The virus-infected tamarillo plants exhibit symptoms of mosaic, blistering, vein-banding, necrotic rings, leaf deformation, and etching in fruits. The virus is transmitted by nematode vectors in a nonpersistent manner. The virus is transmissible by mechanical sap-inoculation, and also by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Solanum betaceum was reported from Colombia, Ecuador, and New Zealand (Eagles et al. 1994; Jaramillo et al. 2011; Sivaprasad et al. 2016b). The virus-infected tamarillo plants exhibit symptoms of mosaic, blistering, vein-banding, necrotic rings, leaf deformation, and etching in fruits. The virus is transmitted by thrips vectors in a persistent propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Cypripedium spp.

811

References Bhargava KS, Joshi RD (1959) A virus disease of tree tomato, Cyphomandra betacea Sendt. due to Potato virus Y. Am J Potato Res 36:8 Chamberlain EE (1954) Plant virus diseases in New Zealand. N Z Dep Sci Ind Res Bull 108:54–56 Eagles R (1994) Tamarillo mosaic potyvirus: characterization and resistance. Thesis PhD, Plant Sciences, School of Biological Sciences. University of Auckland Eagles RM, Gardner RC, Forster RLS (1990) Nucleotide sequence of the Tamarillo mosaic virus coat protein gene. Nucleic Acids Res 18:7166 Eagles R, Gardner R, Forster R (1994) Incidence and distribution of six viruses infecting Tamarillo (Cyphomandra betacea) in New Zealand. N Z J Crop Hortic Sci 22:453–458 Fletcher JD (1987) New plant disease records in New Zealand: additional hosts of alfalfa mosaic virus and cucumber mosaic virus. N Z J Agric Res 30:505–506 Garcia YG, Montoya MM, Gutiérrez PA (2011) Obtención de anticuerpos específicos para la detección del Tamarillo leaf malformation virus (TaLMV) en tomate de árbol. Acta Biol Colomb 16:135–147 Gutierrez PA, Alzate JF, Montoya MM (2015) Genome sequence of a virus isolate from tamarillo (Solanum betaceum) in Colombia: evidence for a new potyvirus. Arch Virol 160:557–560 Insuati ML, Ochoa JB, Martin RR, Alvarez RA, Quito-Avila DF (2016) First report of Potato virus V and Peru tomato mosaic virus on tamarillo (Solanum betaceum) orchards of Ecuador. Plant Dis 100:868 Jaramillo M, Gutierrez PA, Lagos LE, Cotes JM, Marin M (2011) Detection of a complex of viruses in tamarillo (Solanum betaceum) orchards in the Andean region of Colombia. Trop Plant Pathol 36:150–159 Kassanis B, Govier DA (1971) New evidence on the mechanism of aphid transmission of Potato C and Potato aucuba mosaic viruses. J GenVirol 10:99–101 Mossop DW (1977) Isolation, purification and properties of Tamarillo mosaic virus, a member of the Potato virus Y group. N Z J Agric Res 20:535–541 Mossop DW (1982) Potato aucuba mosaic virus – a latent virus of tamarillo (Cyphomandra betacea (Cav.) Sendt). N Z J Agric Res 25:449–453 Sivaprasad Y, Viera W, Patricia G, Orbe K (2015) First report of Potato virus y in tree tomato in Ecuador. J Plant Pathol 97: S73 Sivaprasad Y, Viera W, Buitron J, Orbe K, Ayala L (2016a) First report of Potato leaf roll virus in tree tomato in Ecuador. J Plant Pathol 98:182 Sivaprasad Y, Viera W, Garrido P, Insuasti M (2016b) First report of Tomato spotted wilt virus infecting tree tomato (Solanum betaceum cav.) in Ecuador. J Plant Pathol 98:691 Thomas W, Procter CH (1972) Arabis mosaic virus in Cyphomandra betaceae Sendt. N Z J Agric Res 15:395–404 Villegas MD, Montoya MM, Gutierrez PA (2017) Genome comparison and primer design for detection of Tamarillo leaf malformation virus (TaLMV). Arch Phytopathol Plant Protect 50:713–726 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J GenVirol 98:352–354

Cypripedium spp. Family: Orchidaceae

Cypripedium virus Y Taxonomic position Genus: Potyvirus

Ornamental

(CypVY)

Family: Potyviridae

Geographical distribution CypVY infection in plants of Cypripedium parviflorum and C. franchetii was reported from the UK. It is not clear whether CypVY (recognized by ICTV) is related to a potyvirus from C. calceolus and serologically related to Turnip mosaic virus (Lesemann and Vetten 1985) and later described as

C

812

Cyrtomium falcatum (Japanese holly fern)

Cypripedium calceolus potyvirus and possibly synonymous with Cypripedium chlorotic streak virus, a name which describes the symptoms reported by Lesemann and Vetten (1985). Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically saptransmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, 750 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA. Partial genomic sequence of 1642 nt is available (AF185954) (Gibbs et al. 2000; Revers and Garcia 2015; Wylie et al. 2017).

References Gibbs A, Mackenzie A, Blanchfield AL, Cross P, Wilson CR, Kitajima E, Nightingale M, Clements M (2000) Viruses of orchids in Australia; their identification, biology and control, Australian Orchid Review, vol 65. AOR Publisher, Graphic World, Lewisham, p 3 Lesemann D-E, Vetten HJ (1985) The occurrence of Tobacco rattle and Turnip mosaic viruses in Orchis ssp., and of an unidentified potyvirus in Cypripedium calceolus. Acta Hortic 164:45–54 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J GEN VIROL 98:352–354

Cyrtomium falcatum (Japanese holly fern) Family: Dryopteridaceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Cyrtomium falcatum was reported from the Czech Republic (Mertelik et al. 2002). The virus-infected Japanese holly fern plants exhibit symptoms of concentric necrotic rings and line patterns on the leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Japanese holly fern mottle virus

(JHFMoV)

Taxonomic position JHFMoV is a tentative member of the proposed genus Pteridovirus and family unassigned. Geographical distribution JHFMoV infection is reported in plants of Cyrtomium falcatum from Southern USA (Valverde and Sabanadzovic 2009).

Cyrtosperma spp.

813

Symptoms and host(s) The virus-affected Japanese holly fern plants show various symptoms including mosaic, ringspots, oak leaf patterns, and necrosis; some infected plants showed growth reduction and premature senescence. Transmission The virus is transmitted to healthy Japanese holly fern by grafting, but not by mechanical transmission. Some plants grown from spores collected from infected plants developed symptoms, and JHFMoV was detected, whereas no transmission or symptoms occurred in plants grown from spores of healthy plants (Valverde and Sabanadzovic 2009). Virion properties and genome The virions are quasi-spherical that ranged from 30 to 40 nm in diameter. The genome is a doublestranded RNA. Five double-stranded RNA species were detected and shown to correspond to two genomic RNAs of 6.23 (NC_013133) and 3.01 kb (NC_013134) and three subgenomic RNAs. RNA 1 of the new virus had greatest similarity to members of the genus Idaeovirus, but RNA 2 was shown to have a distinct organization and evolutionary origin, and a new genus “Pteridovirus” was proposed to accommodate JHFMoV (Valverde and Sabanadzovic 2009).

References Mertelik J, Mokra V, Gotzova B, Gabrielova S (2002) Occurrence and identification of Impatiens necrotic spot tospovirus in the Czech Republic. Acta Hortic 568:79–83 Valverde RA, Sabanadzovic S (2009) A novel plant virus with unique properties infecting Japanese holly fern. J Gen Virol 90:2542–2549

Cyrtosperma spp. Family: Araceae

Dasheen mosaic virus Taxonomic position Genus: Potyvirus

Ornamental

(DsMV)

Family: Potyviridae

DsMV infection in plants of Cyrtosperma spp. was reported from Hawaii (Nelson 2008). The virusinfected cyrtosperma plants exhibit systemic mosaic symptoms. The virus is aphid transmitted in a nonpersistent manner, and also through mechanical inoculation. For more details of DsMV, refer to Colocasia esculenta.

References Nelson SC (2008) Dasheen mosaic of edible and ornamental aroids. Plant Dis CTAHR. Available at: http://www.ctahr. hawaii.edu/oc/freepubs/pdf/PD-44.pdf

C

D

Dactylis glomerata (Cocksfoot) Family: Poaceae

Forage crop

Brome mosaic virus Taxonomic position Genus: Bromovirus

(BMV)

Family: Bromoviridae

BMV-infected plants of Dactylis glomerata were reported from Lithuania (Urbanaviciene and Zizyte 2012). The virus-infected cocksfoot plants exhibit mosaic, chlorotic mottling, and streaks on leaves and stem. The virus is transmitted by beetle vectors and is also transmissible by mechanical sap-inoculation. For more details of BMV, refer to Bromus spp.

Cereal yellow dwarf virus RPV Taxonomic position Genus: Polerovirus

(CYDV-RPV)

Family: Luteoviridae

CYDV-RPV infection in plants of Dactylis glomerata was reported from Latvia (Bisnieks et al. 2006). The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of CYDV-RPV, refer to Hordeum vulgare.

Cocksfoot mild mosaic virus Taxonomic position Genus: Panicovirus

(CMMV)

Family: Tombusviridae

© Springer Nature India Private Limited 2019 K. S. Sastry et al., Encyclopedia of Plant Viruses and Viroids, https://doi.org/10.1007/978-81-322-3912-3

816

Dactylis glomerata (Cocksfoot)

Geographical distribution CMMV infection was first reported in plants of Dactylis glomerata from the UK and Germany by Carr (1966) and Huth (1968). The virus spreads in Europe, Canada, the former Czechoslovakia, Denmark, France, Germany, and Norway (Torrance and Harrison 1981; Huth 1988). Symptoms and host(s) The virus-infected cocksfoot plants exhibit faint leaf chlorosis symptoms. The natural host range of this virus includes several plant species of grasses in the genera Bromus, Dactylis, Festuca, Holcus, Phleum and also wheat. Transmission The virus is transmitted by the aphid vector, Myzus persicae (Chamberlain and Catherall 1976). The phleum isolate of this virus is transmitted by the beetles Oulema melanopus and O. lichenis. The virus is transmissible by mechanical sap-inoculation to less than three families; experimentally infected plants mostly show light green streaks. The virus is not transmissible by contact between plants and also not transmitted by seed and pollen. Virion properties and genome The virions are isometric, 28 nm in diameter, and have T = 3 icosahedral symmetry. The genome is a monopartite, linear, single-stranded positive-sense RNA of 4198 nt (EU081018 = NC_011108) (Ziegler 2009; Scholthof 2011).

Cocksfoot mottle virus Taxonomic position Genus: Sobemovirus

(CfMV)

Family: Solemoviridae

Geographical distribution CfMV infection in plants of Dactylis glomerata was first reported from the UK by Serjeant (1964). The virus spreads in Russia, Europe, Canada, France, Germany, Japan, New Zealand, Poland, and the USA (Catherall et al. 1977; Mohamed 1980; Mohamed and Mossop 1981; Toriyama 1982; Lapierre and Hariri 1988; Tamm 1994; Ryabov et al. 1996; Campbell and Guy 2001; Guy 2014; Alderman et al. 2016; Trzmiel and Jezewska 2017). Symptoms and host(s) The virus-infected cocksfoot plants exhibit symptoms of conspicuous yellow streaking and mottling of leaves. The natural host range is narrow, being restricted to a few species and genera in the family Gramineae (Serjeant 1967; Catherall et al. 1977). Transmission The virus is transmitted by chrysomelid beetles, Oulema melanopa and O. lichenis, in a semi-persistent manner. The virus is transmissible by mechanical sap-inoculation. The virus is not transmitted by seed (Serjeant 1967).

Dactylis glomerata (Cocksfoot)

817

Virion properties and genome The virions are isometric, non-enveloped, and 30 nm in diameter (Serjeant 1967). The genome is a single polycistronic positive-sense single-stranded RNA (ssRNA) of 4082 nt (Z48630 = NC_002618; L40905). The genome consists of two overlapping ORFs, ORF2a and ORF2b (Hull 1988; Makinen et al. 1995; Ryabov et al. 1996; Somera et al. 2015).

Cocksfoot streak virus Taxonomic position Genus: Potyvirus

(CSV)

Family: Potyviridae

Geographical distribution CSV infection was first reported in plants of Dactylis glomerata from England by Smith (1952) and Storey (1952). The virus spreads in Europe, Canada, England, France, Holland, Germany, Denmark, Sweden, and the USA (Slykhuis 1958; Smith 1972; Torrance et al. 1994; Urbanaviciene 2001, 2004; Gotz and Maiss 2002). Symptoms and host(s) The virus-infected cocksfoot plants exhibit symptoms of chlorotic streaks on leaves and decreased tillering. Only plants in the Gramineae are susceptible to CSV. In addition to cocksfoot, other natural hosts are Lolium perenne, L. multiflorum, Dactylis polygama, Cynosurus cristatus, and Festuca gigantea. Transmission The virus is transmitted by the aphid vectors, Hyalopteroides humulus, Macrosiphum avenae, M. euphorbiae, Acyrthosiphon dirhodum, and Myzus persicae, in a non-persistent manner (Chamberlain and Catherall 1976). The virus is transmissible by mechanical sap-inoculation to less than three families. The virus is not transmitted by seed. Virion properties and genome The virions are non-enveloped, flexuous filaments, and with a clear modal length of 750 nm and 13 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 9663 nt (AF499738 = NC_003742) (Gotz et al. 1999; Gotz and Maiss 2002; Revers and Garcia 2015; Wylie et al. 2017).

Ryegrass mosaic virus Taxonomic position Genus: Rymovirus

(RGMV)

Family: Potyviridae

RGMV infection in plants of Dactylis glomerata is cosmopolitan in distribution (Torrance et al. 1994). The virus is transmitted by the mite vector Abacarus hystrix in a semi-persistent manner, and is also transmissible through mechanical sap-inoculation. For more details of RGMV, refer to Lolium spp.

D

818

Ryegrass mottle virus Taxonomic position Genus: Sobemovirus

Dactylis glomerata (Cocksfoot)

(RGMoV)

Family: Solemoviridae

RGMoV infection in plants of Dactylis glomerata was reported from Austria, Japan, and Germany (Rabenstein and Huss 2013). The virus-infected cocksfoot plants exhibit symptoms of chlorotic streaks and spots. There is no known vector for this virus. The virus is mechanically sap-transmissible. For more details of RGMoV, refer to Lolium spp.

References Alderman SC, Martin RC, Gilmore BS, Hoffman GD, Sullivan CS, Anderson NP (2016) First report of cocks foot mottle virus infecting Dactylis glomerata in Oregon and the United States. Plant Dis 100:1030 Bisnieks M, Kvarnheden A, Turka I, Sigvald R (2006) Occurrence of barley yellow dwarf virus and cereal yellow dwarf virus in pasture grasses and spring cereals in Latvia. Acta Agric Scand B-Soil Plant Sci 56:171–178 Campbell AW, Guy PL (2001) Cocksfoot mottle virus spreads to the South Island of New Zealand. Aust Plant Pathol 30:217–220 Carr AJH (1966) Report Welsh Plant Breeding Station for 1965, p 90 Catherall PL, Andrews EA, Chamberlain JA (1977) Host-ranges of cocksfoot mottle and cynosurus mottle viruses. Ann Appl Biol 87:233–235 Chamberlain JA, Catherall PL (1976) Association of cocksfoot mild mosaic and cocksfoot streak viruses in Dactylis glomerata and their simultaneous transmission by aphids. Ann Appl Biol 83:475–477 Gotz R, Maiss E (2002) The complete sequence of the genome of Cocksfoot streak virus (CSV), a grass infecting Potyvirus. Arch Virol 147:1573–1583 Gotz R, Huth W, Maiss E (1999) Molecular analysis of the genome of Cockfoot streak potyvirus. J Plant Dis Protect 106:411–417 Guy PL (2014) Viruses of New Zealand pasture grasses and legumes: a review. Crop and Pasture Science 65:841–853 Hull T (1988) The sobemovirus group. In: Koenig R (ed) The plant viruses. Polyhedral virions with monopartite RNA genomes, vol 3. Plenum Press, New York, pp 113–146 Huth W (1968) Untersuchungen über ein neues virus von dactylis glomerata: cocksfoot mild mosaic virus. J Phytopathol 62:300–303 Huth W (1988) Cocksfoot mild mosaic virus (CMMV). In: Smith IM et al (eds) European hand Pl Dis. Blackwell, Oxford, p 70 Lapierre H, Hariri D (1988) Cocksfoot mottle virus. In: Smith IM et al (eds) European hand Pl Dis. Blackwell, London, pp 79–80 Makinen K, Tamm T, Naess V, Truve E, Puurand U, Munthe T, Saarma M (1995) Characterization of cocksfoot mottle sobemovirus genomic RNA and sequence comparison with related viruses. J Gen Virol 76:2817–2825 Mohamed NA (1980) Cocksfoot mottle virus in New Zealand. N Z J Agric Res 23:273–275 Mohamed NA, Mossop DW (1981) Cynosurus and cocksfoot mottle viruses: a comparison. J Gen Virol 55:63 Rabenstein, F., Huss, H. (2013) Studies on grass viruses in Austria. In: 63. Jahrestagung der Vereinigung der Pflanzenzüchter und Saatgutkaufleute Österreichs, Raumberg-Gumpenstein, Österreich. pp 15–18 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Ryabov EV, Krutov AA, Novikov VK, Zheleznikova OV, Yu MS, Zavriev SK (1996) Nucleotide sequence of RNA from the sobemovirus found in infected cocksfoot shows a luteovirus-like arrangement of the putative replicase and protease genes. Phytopathology 86:391–397 Scholthof K-BG (2011) Panicovirus. Tombusviridae. In: The Springer index of viruses. Springer, New York, pp 1911–1916. https://doi.org/10.1007/978-0-387-95919-1_313 Serjeant FP (1964) Cocksfoot mottle virus. Plant Pathol 13:23–24 Serjeant EP (1967) Some properties of cocksfoot mottle virus. Ann Appl Biol 59:31–38 Slykhuis JT (1958) A survey of virus diseases of grasses in northern Europe. FAO Plant Protect Bull 6:129–134 Smith KM (1952) Some recent work on plant viruses. Scient Hortic 11:98–103 Smith KM (1972) A text book of plant virus diseases, 3rd edn. Academic, New York, 684 pp Somera M, Sarmiento C, Truve E (2015) Overview on Sobemoviruses and a proposal for the creation of the family Sobemoviridae. Viruses 7:3076–3115

Dahlia spp.

819

Storey IF (1952) Mosaic on cocksfoot. Plant Pathology 1:101 Tamm T (1994) Characterization of cocksfoot mottle virus. MSc Thesis, Tartu University Toriyama S (1982) Cocksfoot mottle virus in Japan (JE). Ann Phytopathol Soc Jpn 48(4):514–520 Torrance L, Harrison BD (1981) Properties of Scottish isolates of cocksfoot mild mosaic virus and their comparison with others. Ann Appl Biol 97:285–295 Torrance L, Jones AT, Duncan GH (1994) Properties of cocksfoot streak and cocksfoot cryptic, two viruses infecting cocksfoot (Dactylis glomerata) in Scotland. Ann Appl Biol 124:267–281 Trzmiel K, Jezewska M (2017) First report of cocksfoot mottle virus infecting Dactylis glomerata in Poland. Plant Dis 101:1067 Urbanaviciene L (2001) Cocksfoot streak virus in Lithuania. Biologija Nr 4:49–51 Urbanaviciene L, Zizyte M (2012) Identification of Brome mosaic virus in cocksfoot (Dactylis glomerata L.) and meadow fescue (Festuca pratensis Huds.) in Lithuania. Zemdirbyste (Agric) 99:167–172 Urbanaviciene L (2004) Viral diseases of Poaceae family plants. Pemes Ukio Mokslai Nr 1:19–23 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354 Ziegler A, Cowan G, Torrance L (2009) Comparative sequence analysis and serological and infectivity studies indicate that cocksfoot mild mosaic virus is a member of the genus Panicovirus. Arch Virol 154:1545–1549

Dactylorhiza foliosa (Madeiran orchid) Family: Orchidaceae

Ornamental

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Dactylorhiza foliosa was reported from the UK (Skelton et al. 2007). The virus-infected Madeiran orchid plants exhibit chlorotic mottle and streaking symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of BYMV, refer to Phaseolus vulgaris.

References Skelton A, Daly M, Nixon T, Harju V, Mumford RA (2007) First record of Bean yellow mosaic virus infecting a member of the orchid genus Dactylorhiza. Plant Pathol 56:344

Dahlia spp. Family: Asteraceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

Family: Secoviridae

D

820

Dahlia spp.

ArMV infection in plants of Dahlia spp. was reported from Iran and Lithuania (Samuitien_e et al. 2008; Ghotbi and Shahraeen 2005, 2012). The virus-infected dahlia plants were slightly distorted, with symptoms of vein-clearing. Flowers were smaller than normal. The virus is transmitted by the nematode vectors, Xiphinema diversicaudatum and X. coxi, in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Chrysanthemum stunt viroid Taxonomic position Genus: Pospiviroid

(CSVd)

Family: Pospiviroidae

CSVd infection in plants of Dahlia spp. was reported from Japan (Nakashima et al. 2007; Asano et al. 2015). The viroid-infected dahlia plants exhibit symptoms of stunting, small leaves and flowers, and sudden death. The viroid is mechanically sap-transmissible. For more details of CSVd, refer to Chrysanthemum spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Dahlia spp. was reported from New Zealand, Lithuania, Iran, and Bangladesh (Fletcher 1987; Chen and Li 1990; Samuitiene and Navalinskiene 2008; Ara et al. 2012; Ghotbi and Shahraeen 2012). The virus-infected dahlia plants produce a light mosaic with an extreme narrowing of the leaf (fern leaf), and in some cultivars this virus causes oak leaf line patterns across the middle of the leaf (Lawson 1966; Loebenstein et al. 1995). The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Dahlia common mosaic virus

(DCMV)

Taxonomic position DCMV is a tentative member of the genus Caulimovirus and family Caulimoviridae Geographical distribution DCMV-infected plants of Dahlia variabilis were reported from New Zealand, the USA, and the Netherlands. It is probably ubiquitous wherever dahlia is grown (Pappu et al. 2008b; Eid et al. 2009; Hadfield et al. 2011; Chao and Chen 2012; Eid and Pappu 2014). Symptoms and host(s) The virus is only known to infect Dahlia spp., causing variable mosaic and vein-banding symptoms. Transmission The virus is transmitted by aphid vectors. The virus is also mechanically sap-transmissible.

Dahlia spp.

821

Virion properties and genome The virions are isometric, non-enveloped, and 48–50 nm in diameter. The genome is double-stranded DNA, of c. 7950 bp (JN032736) (Hadfield et al. 2011).

Dahlia endogenous plant pararetroviral sequence

(DvEPRS)

Synonyms DMV-D10

D

Taxonomic position DvEPRS is a tentative member of the order Ortervirales Geographical distribution Originally described as DMV-D10, DvEPRS was originally identified in the USA from the cultivated Dahlia variabilis and has also been found in New Zealand, Lithuania, and Egypt, as well as in wild dahlia species growing in their natural habitats in Mexico (Almeyda et al. 2014). DvEPRS is probably ubiquitous where dahlia is grown (Abdel-Salam et al. 2010; Eid et al. 2011). Symptoms and host(s) Currently only known in Dahlia spp., in which transmission occurs vertically through seed, the sequence is only known to be integrated. Virion properties and genome No particles are reported (Pahalawatta et al. 2008). The endogenous (integrated DNA) sequences from different cultivars and species differ considerably, without obvious correlation with geographic origins; the genome of DvEPRS lacks the aphid transmission factor found in typical caulimoviruses such as DMV and DCMV. Full genomic sequences from the USA, New Zealand, and Mexico share identities of 71–97%, with genome lengths of between 7133 and 7159 nt (KC845546; KC967481; KC877995) (Almeyda et al. 2014).

Dahlia latent viroid Taxonomic position Genus: Hostuviroid

(DLVd)

Family: Pospiviroidae

Geographical distribution DLVd was detected in asymptomatic Dahlia spp. with high frequency in Europe, Turkey, and Japan (Verhoeven et al. 2013; Tsushima et al. 2015; Onelge et al. 2017). Symptoms and host(s) The viroid is symptomless in a variety of Dahlia cultivars. Dahlia is the only known host for DLVd, suggesting that the host range of DLVd is narrow. Transmission The viroid is transmissible from Dahlia to Dahlia mechanically and vegetatively.

822

Dahlia spp.

Etiology and genome properties The genome consists of a single-stranded circular RNA of 342 nt (JX263426 = NC_020160), which displays characteristic features of the family Pospiviroidae, i.e., a predicted rodlike secondary structure of minimum free energy with a central conserved region and the ability to form the metastable structure hairpins I and II (Verhoeven et al. 2013). Its central conserved region is identical to that of HpSVd in the genus Hostuviroid; however, DLVd has the terminal conserved region (TCR) characteristic to members of the genus Pospiviroid, but absent in HpSVd. DLVd also lacks the terminal conserved hairpin present in HpSVd. Phylogenetic analysis indicates that HpSVd (genus Hostuviroid) and Pepper chat fruit viroid (genus Pospiviroid) are the closest relatives of DLVd, but DLVd differs from these viroids in its host range, restricted to Dahlia spp. so far (Verhoeven et al. 2013). It has now been assigned to the genus Hostuviroid (Gora-Sochacka 2004; Giguere et al. 2014).

Dahlia mosaic virus Taxonomic position Genus: Caulimovirus

(DMV)

Family: Caulimoviridae

Geographical distribution DMV was first reported in plants of Dahlia pinnata from Germany by Brandenburg (1928). DMV is worldwide in distribution and occurs wherever dahlias are grown (Navalinskiene and Samuitiene 2004; Pappu et al. 2005, 2008a; Pahalawatta et al. 2008; Eid and Pappu 2014; Asano et al. 2015). Symptoms and host(s) The most characteristic symptoms of virus-infected dahlia plants are vein-banding, chlorotic bands adjacent to the midrib, or larger veins of the leaves followed by mosaic and leaf distortion. Shortening of the internodes is also very common in susceptible cultivars giving a short bushy appearance. Other symptoms include chlorosis along the veins which may spread to the entire leaf, and with age, the entire foliage of an infected plant may show severe chlorosis. No change is observed in the color of the petals, but a slight decrease in the number of blooms can be observed in diseased plants. The natural host range of this virus was thought to be limited to Dahlia spp. until a report of infection of chili pepper (Capsicum annuum) in China. Transmission The virus is transmitted by 16 aphid species, including Aphis fabae, Myzus persicae, and Macrosiphum euphorbiae, in a semi-persistent manner. The virus is transmissible by mechanical sap-inoculation, and other members of the Compositae, Solanaceae, Chenopodaeceae, and Amarantheceae can be infected when sap inoculated (Brunt 1971). Pahalawatta et al. (2007) have established that DMV is seedtransmitted through true seed of D. pinnata. The virus was detected in cotyledons and rarely in the seed coat. The virus was also detected in pollen collected from infected plants (Pappu et al. 2005). The primary spread of the virus takes place through the use of virus-infected vegetative planting material. Virion properties and genome The virions are isometric, non-enveloped, and 48–50 nm in diameter with no obvious surface structure. The genome is a monopartite, circular double-stranded DNA of c. 7900 bp (NC_018616) (Nicolaisen 2003; Hohn 2011; Sahoo et al. 2015).

Dahlia spp.

823

Groundnut bud necrosis orthotospovirus Taxonomic position Genus: Tospovirus

(GBNV)

Family: Peribunyaviridae

GBNV infection in plants of Dahlia coccinea was reported from India (Holkar et al. 2013). The virusinfected dahlia plants showed mottle, distortion, and marginal chlorosis of the leaves, in addition to bud necrosis. The virus is transmitted by thrips vector in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of GBNV, refer to Arachis hypogaea.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Tospovirus

(INSV)

Family: Peribunyaviridae

INSV infection in plants of Dahlia spp. was reported from the USA, Italy, and Iran (Roggero et al. 1999; Ghotbi et al. 2005). The virus-infected dahlia plants exhibit foliar spots and ringspot symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Mungbean yellow mosaic virus Taxonomic position Genus: Begomovirus

(MYMV)

Family: Geminiviridae

MYMV infection in plants of dahlia hybrid was reported from Bangladesh (Ara et al. 2002). The virusinfected dahlia plants exhibit yellow mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of MYMV, refer to Vigna radiata.

Potato spindle tuber viroid Taxonomic position Genus: Pospiviroid

(PSTVd)

Family: Pospiviroidae

PSTVd isolate was detected from Dahlia spp. in Japan and in the Netherlands (Tsushima et al. 2011, 2015; Pest report-The Netherlands 2013; IPPC 2014; Verhoeven et al. 2016). No symptoms appeared in dahlia cultivars in the first season of infection; however, the growth of infected plants was reduced, and tubers of infected plants were small, spindle-shaped, or elongated in the following season (Fujiwara et al. 2013). The viroid is transmissible mechanically to tomato plants (Tsushima et al. 2011). For more details of PSTVd, refer to Solanum tuberosum.

D

824

Dahlia spp.

Potato virus Y

(PVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

PVY infection in plants of Dahlia spp. is cosmopolitan in distribution (Raj et al. 1999). The virusinfected Dahlia pinnata plants exhibit severe mosaic on newly emerging leaves followed by extreme stunting and deformed flowers. The virus is aphid transmitted in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of PVY, refer to Solanum tuberosum.

Tobacco mosaic virus

(TMV)

Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

TMV infection in plants of Dahlia variabilis was reported from India (Reddy and Sumanwar 1991). There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

The first detection of TSV in the UK was in 5 of 73 cultivars of dahlia tested; three cultivars infected only with TSV were symptomless, while inoculation of seedlings yielded only indistinct symptoms within 6 months (Brunt 1968). TSV has also been detected infecting a few plants of Dahlia variabilis in New Zealand (Pappu et al. 2008a). TSV is widespread in dahlia (Mokra et al. 2008; Harju et al. 2011). The virus-infected dahlia plants are often symptomless or express inconspicuous leaf symptoms or light mottling. The petals also exhibit discoloration and deformation (flower breaking). White stripes in blossoms were most frequently expressed in the second half of the flowering season. The virus is transmitted by the thrips vectors (Frankliniella spp. and Thrips tabaci), is present in/on pollen, and enters in to the host through the injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible, but is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Dahlia spp. was reported from Iran (Ghotbi and Shahraeen 2009, 2012). The virus is transmitted by nematode vectors in a non-persistent manner. The virus is also transmissible by mechanical sap-inoculation, and by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Dahlia spp.

825

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Tospovirus

(TSWV)

Family: Peribunyaviridae

TSWV was first described as “oak leaf disease” or “dahlia ring spot” based on symptoms (Brierley 1933a, b), and subsequently this virus infection in dahlia plants was reported from the USA, Japan, Iran, and the Netherlands (Inouye and Inouye 1973; Daughtrey 1996; Derks and Lemmers 1996; van Schadewijk 1996; Daughtrey et al. 1997; Ghotbi and Shahraeen 2012; Asano et al. 2015). The virusinfected dahlia plants exhibit different types of ringspot symptoms. Another typical symptom is chlorotic lines on the leaves which sometimes become necrotic. In some cultivars several irregular and sinuous lines or concentric rings are produced on older leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow ring virus

(TYRV)

Taxonomic position TYRV is a tentative member of the genus Tospovirus and family Peribunyaviridae TYRV was reported from Iran affecting Dahlia spp. (Ghotbi and Shahraeen 2012). The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of TYRV, refer to Solanum lycopersicum.

References Ara MR, Masud MMH, Akanda AM (2012) Detection of plant viruses in some ornamental plants that act as alternate hosts. Agriculturists 10:46–54 Asano S, Matsushita Y, Hirayama Y, Naka T (2015) Simultaneous detection of tomato spotted wilt virus, Dahlia mosaic virus and chrysanthemum stunt viroid by multiplex RT-PCR in dahlias and their distribution in Japanese dahlias. Lett Appl Microbiol 61(2):113–120 Brandenburg E (1928) Uber mosaikkrankheiten an Compisite. Forsch Gebeit Pflanzenkrankh Immunität Pflanzenreich 5:39–72 Brierley P (1933a) Dahlia mosaic and its relationship to stunt. Am Dahlia Soc Ser 9:6–11 Brierley P (1933b) Studies on mosaic and related diseases on dahlia. Boyce Thompson Inst Contrib 5:235–288 Brunt AA (1968) Tobacco streak virus in dahlias. Plant Pathol 17:119–122 Brunt AA (1971) Some hosts and properties of Dahlia mosaic virus. Ann Appl Biol 67:357–368 Chao HY, Chen YK (2012) Molecular characterization of a dahlia mosaic-associated Caulimovirus. Plant Pathol Bull 21:110–114 Chen JH, Li DB (1990) A preliminary study of a potyvirus and cucumber mosaic virus infecting Dahlia pinnata. Acta Agric Univ Zhejiangensis 16(Suppl. 2):78–82 Daughtrey ML (1996) Detection and identification of tospoviruses in greenhouses. Acta Hortic 431:90–98 Daughtrey ML, Jones RK, Moyer JW, Daub ME, Baker JR (1997) Tospoviruses strike the greenhouse industry; INSV has become a major pathogen on flower crops. Plant Dis 81:1220–1230 Derks AFLM, Lemmers MEC (1996) Detection of tospoviruses in bulbous crops and transmissibility by vegetative propagation. Acta Hortic 432:132–137 Eid S, Pappu HR (2014) Biological studies of three caulimoviruses associated with dahlia (Dahlia variabilis). Can J Plant Pathol 36:110–115

D

826

Dahlia spp.

Eid S, Druffel KL, Saar DE, Pappu HR (2009) Incidence of multiple and distinct species of Caulimoviruses in Dahlia (Dahlia variabilis). Hortic Sci 44:1498–1500 Fletcher JD (1987) New plant disease records in New Zealand: additional hosts of alfalfa mosaic virus and cucumber mosaic virus. N Z J Agric Res 30:505–506 Fujiwara Y, Nomura Y, Hiwatashi S, Shiki Y, Itto T, Hamanaka D, Saito N (2013) Pathogenicity to potato of Potato spindle tuber viroid isolated from dahlia and its transmissibility in dahlia. Res Bull Plant Protect Jpn 49:41–46 Ghotbi T, Shahraeen N (2005) First report on incidence of arabis mosaic virus (ArMV, Nepovirus) on ornamental plants in Iran. Iran J Plant Pathol 41(2):305–306 Ghotbi T, Shahraeen N (2009) Natural incidence and infectivity level of three nepoviruses in ornamental crops in Iran. J Plant Breed Crop Sci 1:39–44 Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses affecting propagated ornamentals in Northern Iran. Int Res J Microbiol 3:373–381 Ghotbi T, Sharaeen N, Winter S (2005) Occurrence of tospoviruses in ornamental and weed species in Markazi and Tehran provinces in Iran. Plant Dis 89:425–429 Giguere T, Raj Adkar-Purushothama C, Perreault J-P (2014) Comprehensive secondary structure elucidation of four genera of the family pospiviroidae. PLoS One 9(6):e98655. https://doi.org/10.1371/journal.pone.0098655 Gora-Sochacka A (2004) Viroids: unusual small pathogenic RNAs. Acta Biochim Pol 51(3):587–607 Hadfield J, Linderme D, Shepherd DN, Bezuidenhout M, Lefeuvre P, Martin DP, Varsani A (2011) Complete genome sequence of a dahlia common mosaic virus isolate from New Zealand. Arch Virol 156:2297–2301 Harju VA, Skelton AL, Monger WA, Forde SMD, Daly M, Nixon T, Lawson R, Bennett S, Mumford RA (2011) A report of viruses, viroids and phytoplasmas found in ornamental plants from 1999–2007: findings from central science laboratory, UK. Acta Hortic 901:223–229 Hohn T (2011) Caulimovirus. Caulimoviridae. In: The Springer index of viruses. Springer, New York, pp 271–277. https://doi.org/10.1007/978-0-387-95919-1_41 Holkar SK, Mandal B, Jain RK (2013) New ornamental and cucurbitaceous hosts of groundnut bud necrosis virus in India. Indian J Virol 24:146 Inouye T, Inouye N (1973) Tomato spotted wilt virus isolated from Dahlia (studies in the viruses of plants in Compositae in Japan III). Ber Ohara Inst Landwirtsch Biol Okayama Univ 15:25 IPPC (2014) First finding of potato spindle tuber viroid (PSTVd) in Dahlia sp. corm production in the Netherlands, IPPC Official Pest Report, No. NLD-21/1. FAO, Rome Lawson RH (1966) Oak leaf chlorosis symptomatic of cucumber mosaic infection in dahlia. Phytopathology 56:343–344 Loebenstein G, Lawson RH, Brunt AA (1995) Virus and virus-like diseases of bulb and flowering crops. Wiley, Chichester, p 543 Mokra V, Gotzova B, Bezdekova V, Dedic P, Ptacek J (2008) First report of tobacco streak virus on dahlia in the Czech Republic. Plant Dis 92:484 Nakashima A, Hosokawa M, Maeda S, Yazawa S (2007) Natural infection of Chrysanthemum stunt viroid in dahlia plants. J Gen Plant Pathol 73:225–227 Navalinskiene M, Samuitiene M (2004) Virological evaluation of the Lithuanian dahlia (Dahlia Cav.), gladiolus (Gladiolus L.), iris (Iris L.) and peony (Paeonia L.) cultivars and hybrids. Biologija 3:57–63 Nicolaisen M (2003) Partial molecular characterization of Dahlia mosaic virus and its detection by PCR. Plant Dis 87:945–948 Onelge N, Ertugrul B, Guler P (2017) First report of dahlia latent viroid in Turkey. J Plant Pathol 99:812 Pahalawatta V, Druffel K, Pappu HR (2007) Seed transmission of Dahlia mosaic virus in Dahlia pinnata. Plant Dis 91:88–91 Pahalawatta V, Druffel KL, Wyatt SD, Eastwell KC, Pappu HR (2008) Genome structure and organization of a member of a novel and distinct species of the genus Caulimovirus associated with dahlia mosaic. Arch Virol 153:733–738 Pappu HR, Wyatt SD, Druffel KL (2005) Dahlia moaic virus: molecular detection and distribution in dahlia in the US. Hortic Sci 40:697–699 Pappu HR, Hammett KRW, Druffel KL (2008a) Dahlia mosaic virus and Tobacco streak virus in dahlia (Dahlia variabilis) in New Zealand. Plant Dis 92:1138 Pappu HR, Druffel KL, Miglino R, van Schadewijk AR (2008b) Nucleotide sequence and genome organization of a member of a new and distinct Caulimovirus species from dahlia. Arch Virol 153:2145–2148 Pest Report – The Netherlands (2013) First finding of Potato Spindle Tuber Viroid (PSTVd) in Dahlia sp. corm production in the Netherlands, NEPO September 2013 Raj SK, Ainuddin, Abidi SMH, Singh BP (1999) Characterization of a virus causing severe mosaic and floral deformations of Dahlia pinnata L. in India. Indian J Plant Pathol 17:8–12 Reddy OR, Sumanwar AS (1991) Detection of Tobacco mosaic virus in imported dahlias (Dahlia variabilis) from Netherlands. Indian J Plant Protect 19:218–220 Roggero P, Ciuffo M, Dellavalle G, Gotta P, Gallo S, Peters D (1999) Additional ornamental species as hosts of impatiens necrotic spot tospovirus in Italy. Plant Dis 83:967

Daiswa polyphylla

827

Sahoo DK, Sarkar S, Raha S, Das NC, Banerjee J, Maiti IB (2015) Analysis of dahlia mosaic virus full-length transcript promoter-driven gene expression in transgenic plants. Plant Mol Biol Rep 33:178–199 Samuitiene M, Navalinskiene M (2008) Occurrence of cucumber mosaic cucumovirus on ornamental plants in Lithuania. Zemdirbyste-Agric 95:135–143 Samuitien_e M, Navalinskien_e M, Jackevičien_e E (2008) Arabis mosaic virus on ornamental plants. Biologija 54:264–268 Tsushima T, Murakami S, Ito H, He Y-H, Charith Raj AP, Sano T (2011) Molecular characterization of potato spindle tuber viroid in dahlia. J Gen Plant Pathol 77:253–256 Tsushima T, Matsushita Y, Fuji S, Sano T (2015) First report of Dahlia latent viroid and Potato spindle tuber viroid mixedinfection in commercial ornamental dahlia in Japan. New Dis Rep 31:11 van Schadewijk AR (1996) Detection of Tomato spotted wilt virus in dahlia. Acta Hortic (ISHS) 432:384–391 Verhoeven JTHJ, Meekes ETM, Roenhorst JW, Flores R, Serra P (2013) Dahlia latent viroid: a recombinant new species of the family Pospiviroidae posing intriguing questions about its origin and classification. J Gen Virol 94:711–719 Verhoeven JTJ, Westenberg M, van Ede EPM, Visser K, Roenhorst JW (2016) Identification and eradication of potato spindle tuber viroid in dahlia in the Netherlands. Eur J Plant Pathol 146:443–447

Daiswa polyphylla Synonyms Paris polyphylla Family: Melanthiaceae

Paris mosaic necrosis virus (PMNV) Taxonomic position Genus: Potyvirus Family: Potyviridae Geographical distribution PMNV infection in plants of Paris polyphylla var. yunnanensis was reported from southern China (Lan et al. 2017). Symptoms and host(s) The virus-infected plants exhibit mottle, necrotic and mosaic symptoms on leaves. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is transmitted by mechanical sap-inoculation. The virus is not transmitted by contact between plants; not transmitted by seed; not transmitted by pollen. Virion properties and genome The virions are non-enveloped, flexuous filaments 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 9660 nt (MF509898) (Revers and Garcia 2015; Lan et al. 2017) and contains one large ORF encoding a polyprotein of 353 kDa, and includes the nine semi-conserved proteolytic cleavage sites. A short ORF resulting from transcriptional slippage in the P3 cistron generates P3N-PIPO (Wylie et al. 2017).

D

828

Dalechampia spp.

References Lan P, Zhao J, Zhou Y, Li Y, Shen D, Liao Q, Li R, Li F (2017) Complete genome sequence of Paris mosaic necrosis virus, a distinct member of the genus Potyvirus. Arch Virol 163(3):787–790 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: potyviridae. J Gen Virol 98:352–354

Dalechampia spp. Family: Euphorbiaceae

Ornamental

Dalechampia chlorotic mosaic virus Taxonomic position Genus: Begomovirus

(DaChMV)

Family: Geminiviridae

Geographical distribution DaChMV infection in plants of Dalechampia spp. was reported from Venezuela (Fiallo-Olive et al. 2013). Symptoms and host(s) The virus-infected Dalechampia plants exhibit chlorotic mosaic symptoms. The virus has been isolated from Dalechampia and from Boerhavia diffusa (Fiallo-Olive et al. 2013). Transmission The transmission of DaChMV has not been investigated. It is likely that, in common with other begomoviruses, DaChMV is transmitted by the whitefly vector Bemisia tabaci in a circulative, nonpropagative manner. Virion properties and genome The virion structure of DaChMV has not been investigated. In common with all geminiviruses, the virions of DaChMV are likely geminate (twinned quasi-icosahedra). DaChMV is typical of the majority of begomoviruses native to the New World with a bipartite genome consisting of two circular, single-stranded DNA components. DNA-A contains 2616 nt (JN848775 = NC_018716) and DNA-B of 2598 nt (JN848776 = NC_018718) (Briddon 2001; Fiallo-Olive et al. 2013; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the New World, the DNA-A component of DaChMV encodes five genes, one in the virionsense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for DaChMV.

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619

Daphne spp.

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Fiallo-Olive E, Chirinos DT, Geraud-Pouey F, Moriones E, Navas-Castillo J (2013) Complete genome sequences of two begomoviruses infecting weeds in Venezuela. Arch Virol 158(1):277–280 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

D

Daphne spp. Family: Thymelaeaceae

Ornamental

Alfalfa mosaic virus

(AMV)

Taxonomic position Genus: Alfamovirus

Family: Bromoviridae

AMV infection in plants of Daphne odora was reported from the USA and New Zealand (Milbrath and Young 1956; Forster and Milne 1975). The virus-infected daphne plants exhibit mosaic symptoms although some cultivars are symptomless. The virus is transmitted by a large number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

ArMV has been reported from plants of Daphne odora and D. dauphinii in New Zealand and earlier in the UK (Forster and Milne 1975). No distinct symptoms were associated with ArMV infection. The virus is transmitted by the nematode vectors (Xiphinema spp.) in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Carnation mottle virus Taxonomic position Genus: Alphacarmovirus

(CarMV)

Family: Tombusviridae

CarMV was detected in plants of Daphne odora and D. burkwoodii in New Zealand and Korea (MorrisKrsinich and Milne 1977; Lee et al. 2006a). The virus-infected daphne plants exhibit symptomless infections. The virus is transmitted by means not involving a vector. The virus is transmissible by mechanical sap-inoculation, by grafting, and also by contact between plants. For more details of CarMV, refer to Dianthus caryophyllus.

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Daphne spp.

Cucumber mosaic virus

(CMV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV infection in plants of Daphne spp. was reported from the USA, New Zealand, the UK, and Korea (Smith 1952; Milbrath and Young 1956; Forster and Milne 1975; Lee et al. 2006a; Lee and Ryu 2009). The virus-infected daphne plants are either symptomless or show mosaic symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Cycas necrotic stunt virus Taxonomic position Genus: Nepovirus

(CNSV)

Family: Secoviridae

CNSV infection in plants of Daphne odora was reported from Korea (Lee et al. 2006a; Lee and Ryu 2006, 2009). The virus-infected daphne plants exhibit chlorosis and stunting symptoms. The virus is mechanically sap-transmissible. For more details of CNSV, refer to Cycas spp.

Daphne mosaic virus Taxonomic position Genus: Potyvirus

(DapMV)

Family: Potyviridae

Geographical distribution DapMV infection in plants of Daphne mezereum was reported from the Czech Republic (Franova et al. 2006).

Symptoms and host(s) The leaves of infected daphne plants exhibit light green rings and mosaic symptoms.

Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically saptransmissible.

Virion properties and genome The virions are non-enveloped, flexuous filaments of c.696 nm length and 13 nm width. The genome consists of a single molecule of linear, positive-sense ssRNA of 9548 nt (DQ299908 = NC_008028) (Petrzik and Franova 2006; Revers and Garcia 2015).

Daphne spp.

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Daphne virus S

(DVS)

Taxonomic position Genus: Carlavirus

Family: Betaflexiviridae

Geographical distribution DVS was first reported in plants of Daphne spp. from New Zealand by Forster and Milne (1978). The virus spreads in New Zealand and Korea (Lee et al. 2006a; Lee and Ryu 2009). Symptoms and host(s) The virus appeared to be latent in most Daphne species and cultivars but was associated with foliar symptoms in D. odora Thunb. ‘Leucanthe’ (the most important cultivar in New Zealand). The virusinfected D. odora ‘Leucanthe’ specimens growing in unshaded sites displayed leaf distortion and downcurling of leaf edges, particularly in mature leaves. Transmission The virus is transmitted by the aphid vector, Myzus persicae, in a non-persistent manner. The virus is not transmissible by contact between plants. Virion properties and genome The virions are flexuous filaments about 720 nm in length and 13 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8739 nt (NC_008020 = AJ620300) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Lee et al. 2003, 2006b; Adams et al. 2004; Wylie et al. 2017).

Daphne virus Y

(DVY)

Synonyms Daphne mottle virus (DapMoV) Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution DVY infection in plants of Daphne spp. was reported from Australia, the Netherlands, New Zealand, Germany, Korea, and the UK (Forster and Milne 1976; Lee et al. 2006a; Lee and Ryu 2009; Igori et al. 2016; Park et al. 2016). Symptoms and host(s) The virus-infected daphne plants exhibit symptoms of mosaic, chlorotic mottle, chlorosis, leaf distortion, and decreased flower quality. Transmission The virus is transmitted by the aphid vector Myzus persicae in a non-persistent manner. The virus is mechanically sap-transmissible.

D

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Daphne spp.

Virion properties and genome The virions are non-enveloped, flexuous filaments of 800 nm long and 12 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 9448 nt (KU556609) (excluding the 30 -terminal poly(A) tail) encoding a polyprotein of 3,065 amino acids, with nine putative protease cleavage sites producing ten proteins (Igori et al. 2016; Wylie et al. 2017).

Tobacco mosaic virus

(TMV)

Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

TMV infection in plants of Daphne odora was reported from New Zealand (Forster and Milne 1975). There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV was detected in a small number of Daphne odora plants in New Zealand (Forster and Milne 1975). The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato bushy stunt virus Taxonomic position Genus: Tombusvirus

(TBSV)

Family: Tombusviridae

TBSV infection in plants of Daphne mezereum was reported from the Czech Republic (Novak and Lanzova 1980). The virus is transmissible by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants. There is no known vector for this virus. For more details of TBSV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Daphne mezereum was reported from Victoria and Australia (Chiko and Godkin 1984). The virus-infected daphne plants exhibit symptoms of mottling of leaves. The virus is

Daphne spp.

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transmitted by nematode vectors in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and also by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV infection in plants of Daphne odora was reported from Korea (Lee and Ryu 2009). The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of WMV, refer to Citrullus lanatus.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Chiko AW, Godkin SE (1984) First report of Tomato ringspot virus infecting daphne. Plant Dis 68:536 Forster RLS, Milne KS (1975) Survey of viruses infecting daphne in New Zealand. NZ J Agric Res 18:391–398 Forster RLS, Milne KS (1976) Daphne virus Y: potyvirus from daphne. N Z J Agric Res 19:359–371 Forster RLS, Milne KS (1978) Daphne virus S: a carlavirus from daphne. N Z J Agric Res 21:131–135 Franova J, Petrzik K, Lesemann DE, Navratil M (2006) Daphne mosaic virus (DapMV), a new potyvirus from Daphne mezereum in the Czech Republic. Arch Virol 151:793–801 Igori D, Hwang US, Lim S, Zhao F, Kwon S-K, Moon JS (2016) The first complete sequence and genome structure of Daphne virus Y. Arch Virol 161:2905–2908 Lee BY, Ryu KH (2006) Identification and molecular detection of Cycas necrotic stunt virus from Daphne odora plants in Korea. Hortic Environ Biotechnol 47:75–79 Lee BY, Ryu KH (2009) Incidence of virus diseases and RT-PCR detection of Daphne-infecting viruses in Korea. Eur J Plant Pathol 124:127–132 Lee BY, Choi SH, Ryu KH (2003) Characterization of the 30 -terminal nucleotide sequence of two Korean isolates of Daphne virus S support its placement as a distinct species of Carlavirus. Arch Virol 148:1915–1924 Lee BY, Ha JH, Ryu KH (2006a) Daphne-infecting viruses in Korea and properties of Daphne virus S. Acta Hortic 722:55–58 Lee BY, Min BE, Ha JH, Lee MY, Paek KH, Ryu KH (2006b) Genome structure and the complete sequence of genome RNA of Daphne virus S. Arch Virol 151:193–200 Milbrath JA, Young RA (1956) Cucumber mosaic virus and alfalfa mosaic virus isolated from Daphne odora. Plant Dis Reptr 40:279–283 Morris-Krsinich BA, Milne KS (1977) Natural infection of Daphne by Carnation mottle virus. Plant Dis Reptr 61:675–678 Novak JB, Lanzova J (1980) Demonstration of tomato bushy stunt virus in some forest tree species and plants. Lesnictví 26:1009–1016 Park CY, Park J, Lee B-J, Bak S, Lee H-K, Kim J-S, Youn Y, Suh SJ, Lee S-H (2016) Identification of Daphne mottle virus isolated from Daphne odora, a new member of the genus Potyvirus. Res Plant Dis 22:59–63 Petrzik K, Franova J (2006) Complete genome sequence of Daphne mosaic virus – a potyvirus from an ornamental shrub related to papaya leaf distortion mosaic virus. Arch Virol 151:1461–1465 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Smith KM (1952) Some garden plants susceptible to infection with the cucumber mosaic virus. J R Hortic Sci 77:19–21 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354

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834

Daphniphyllum teysmannii

Daphniphyllum teysmannii Family: Daphniphyllaceae

Trees/Shrubs

Satsuma dwarf virus

(SDV)

Taxonomic position Genus: Sadwavirus

Family: Secoviridae

SDV was detected in plants of Daphniphyllum teysmannii without any clear symptoms, growing near orchards of satsuma mandarin in Japan (Nakazono-Nagaoka et al. 2014). The virus is transmissible through the soil and is also transmissible by mechanical sap-inoculation. No vector has been identified for this virus. The virus is not transmissible by contact between plants. For more details of SDV, refer to Citrus spp.

References Nakazono-Nagaoka E, Takemoto S, Fujikawa T, Nakajima K, Uenishi H, Iwanami T (2014) Natural Satsuma dwarf infection of two woody plants, Daphniphyllum teijsmannii Zoll. ex Kurz. and Viburnum odoratissimum Ker-Gaul. var. awabuki (K. Koch) Zabel near Citrus Fields. JARQ 48:419–424

Datura spp. (Jimson weed) Family: Solanaceae

Weed host

Chilli veinal mottle virus Taxonomic position Genus: Potyvirus

(ChiVMV)

Family: Potyviridae

ChiVMV infection in plants of Datura inoxia was reported from India (Kaur et al. 2015). The virusinfected datura plants exhibit severe mottling and distortion symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation, and by grafting. For more details of ChiVMV, refer to Capsicum annuum.

Colombian datura virus Taxonomic position Genus: Potyvirus

(CDV)

Family: Potyviridae

Datura spp. (Jimson weed)

835

Geographical distribution CDV was first reported in plants of Datura sanguinea and Datura candida from Colombia by Kahn and Bartels (1968). The virus spreads in Colombia. Symptoms and host(s) The virus-infected Jimsonweed plants exhibit symptoms of vein-banding and chlorotic flecking followed by mottling.

D Transmission The virus is transmitted by an aphid vector, Myzus persicae, in a non-persistent manner. The virus is also mechanically sap-transmissible and by grafting. Virion properties and genome The virions are non-enveloped, flexuous filaments 800 nm long and 12 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 9621 nt (JQ801448 = NC_020072) (Revers and Garcia 2015; Wylie et al. 2017).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Datura innoxia was reported from India (Raj et al. 1999). The virus-infected Jimsonweed plants exhibit green mosaic, leaf deformation, and shoestring symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of CMV, refer to Cucumis sativus.

Datura leaf curl virus

(DaLCV)

Taxonomic position Genus: Begomovirus Family: Geminiviridae Geographical distribution DaLCV infection in plants of Datura inoxia was reported from Sudan (Mohammed et al. 2018). Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2782 nt (MF402918) (Brown et al. 2015; Zerbini et al. 2017; Mohammed et al. 2018).

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Datura spp. (Jimson weed)

Datura leaf distortion virus Taxonomic position Genus: Begomovirus

(DaLDV)

Family: Geminiviridae

Geographical distribution DaLDV infection in plants of Datura stramonium was reported from Venezuela (Fiallo-Olive et al. 2013). Symptoms and host(s) The virus-infected Jimsonweed plants exhibit mosaic and leaf distortion symptoms. Transmission The transmission of the DaLDV has not been investigated. It is likely that, in common with other begomoviruses, DaLDV is transmitted by the whitefly vector Bemisia tabaci in a circulative, nonpropagative manner. Virion properties and genome The virion structure of DaLDV has not been investigated. In common with all geminiviruses, the virions of DaLDV are likely geminate (twinned quasi-icosahedra). DaLDV is typical of the majority of begomoviruses native to the New World with a bipartite genome consisting of two circular, single-stranded DNA components. DNA-A contains 2599 nt (JN848773 = NC_018717) and DNA-B of 2564 nt (JN848774 = NC_018715) (Briddon 2001; Fiallo-Olive et al. 2013; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the New World, the DNA-A component of DaLDVencodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for DaLDV.

Datura shoestring virus Taxonomic position Genus: Potyvirus

(DSSV)

Family: Potyviridae

DSSV infection in plants of Datura metel was reported from Nigeria (Taiwo et al. 2006). The virusinfected Jimsonweed plants exhibit blistering mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of DSSV, refer to Solanum jasminoides.

Datura yellow vein nucleorhabdovirus Taxonomic position Genus: Nucleorhabdovirus

(DYVV)

Family: Rhabdoviridae

Datura spp. (Jimson weed)

837

Geographical distribution DYVV was first reported in plants of Datura stramonium from South East Queensland, Australia, by Thomas and Behncken (1983). The virus spreads in Australia (Thomas and Dietzgen 1991; Dietzgen et al. 2013). Symptoms and host(s) The virus-infected Jimsonweed plants exhibit symptoms of vein yellowing and leaf and flower bud malformation. This virus infects tomato and shows vein-clearing and chlorosis, mottling, and leaf malformation. Transmission The virus is not mechanically sap-transmissible. Aphid species failed to transmit this virus. The virus is not transmissible by contact between plants and also not through seed. The virus is transmissible by grafting. Virion properties and genome The virions are enveloped and bacilliform and measure 77 nm in diameter and 166 nm long. There are distinct surface projections (spikes) dispersed evenly over all the surface. The genome is a monopartite, negative-sense, single-stranded RNA of 13,188 nt (KM823531 = NC_028231) (Jackson et al. 2011; Dietzgen et al. 2015; Walker et al. 2018).

Pepper veinal mottle virus Taxonomic position Genus: Potyvirus

(PVMV)

Family: Potyviridae

PVMV infection in plants of Datura stramonium was reported from Nigeria (Taiwo et al. 2004, 2006). The virus-infected Jimsonweed plants exhibit mosaic and leaf distortion symptoms. This virus is transmitted in a non-persistent manner by several aphid species and is also transmissible by mechanical sap-inoculation. For more details of PVMV, refer to Capsicum annuum.

Potato spindle tuber viroid Taxonomic position Genus: Pospiviroid

(PSTVd)

Family: Pospiviroidae

PSTVd infection in plants of Datura spp. was reported from Australia and the Czech Republic (Mackie et al. 2016; Matousek et al. 2014). The viroid-infected Jimsonweed plants exhibit symptoms of stunting, leaf chlorosis, and/or reduced leaf size. The viroid is mechanically sap-transmissible. For more details of PSTVd, refer to Solanum tuberosum.

Potato virus Y

(PVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

D

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Datura spp. (Jimson weed)

PVY infection in plants of Datura spp. was reported from India (Prasada Rao and Yaraguntaiah 1976; Garg et al. 2003). The virus-infected Jimsonweed plants exhibit mosaic mottling symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation to a large number of hosts and by grafting. For more details of PVY, refer to Solanum tuberosum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Datura spp. was reported from Australia (Sharman et al. 2015). The virusinfected Jimsonweed plants exhibit necrotic spots and ringspot symptoms. The virus is transmitted by the thrips vectors, is present in/on pollen, and enters into the host through the injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible but is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tobacco vein banding mosaic virus Taxonomic position Genus: Potyvirus

(TVBMV)

Family: Potyviridae

TVBMV infection in plants of Datura stramonium was reported from China (Roggero et al. 2000). The virus-infected Jimsonweed plants show severe mosaic with blistering of the leaves. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of TVBMV, refer to Nicotiana tabacum.

Tomato chlorosis virus Taxonomic position Genus: Crinivirus

(ToCV)

Family: Closteroviridae

ToCV infection in plants of Datura stramonium was reported from South Africa (Moodley et al. 2016). The virus-infected Jimsonweed plants exhibit severe interveinal leaf chlorosis symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a semi-persistent manner. For more details of ToCV, refer to Solanum lycopersicum.

Tomato chlorotic spot orthotospovirus Taxonomic position Genus: Tospovirus

(TCSV)

Family: Peribunyaviridae

TCSV infection in plants of Datura stramonium was reported from Puerto Rico (Webster et al. 2013). The virus-infected Jimsonweed plants exhibit ringspot, irregular chlorotic areas, and deformation of

Datura spp. (Jimson weed)

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leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of TCSV, refer to Solanum lycopersicum.

Tomato yellow leaf curl China virus Taxonomic position Genus: Begomovirus

(TYLCCNV)

Family: Geminiviridae

TYLCCNV infection in plants of Datura stramonium was reported from China (Ding et al. 2007). The virus-infected Jimsonweed plants exhibit chlorotic and leaf curl symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not transmissible by contact between plants. For more details of TYLCCNV, refer to Solanum lycopersicum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Datura stramonium was reported from Spain and Cyprus (Jorda et al. 2001; Papayiannis et al. 2011). The virus-infected Jimsonweed plants exhibit symptomless infections. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not transmissible by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

Watermelon chlorotic stunt virus Taxonomic position Genus: Begomovirus

(WmCSV)

Family: Geminiviridae

WmCSV infection in plants of Datura innoxia was reported from Sudan (Mohammed et al. 2017). The virus-infected datura plants exhibit yellowing and leaf deformation symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by mechanical inoculation, but is transmissible by grafting. For more details of WmCSV, refer to Citrullus lanatus.

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Dietzgen RG, Tan ER, Yong AHS, Feng C-WA (2013) Partial polymerase gene sequence, phylogeny and RT-PCR diagnostic assay for Datura yellow vein nucleorhabdovirus. Aust Plant Dis Notes 8:21–25

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Datura spp. (Jimson weed)

Dietzgen RG, Innes DJ, Bejerman N (2015) Complete genome sequence and intracellular protein localization of Datura yellow vein nucleorhabdovirus. Virus Res 205:7–11 Ding M, Luo YQ, Dong JH, Fang Q, Zhang ZK (2007) First report of Tomato yellow leaf curl China virus with DNA b infecting Datura stramonium in China. Aust Plant Dis Notes 2:63 Fiallo-Olive E, Chirinos DT, Geraud-Pouey F, Moriones E, Navas-Castillo J (2013) Complete genome sequences of two begomoviruses infecting weeds in Venezuela. Arch Virol 158(1):277–280 Garg ID, Shiv Kumar, Paul Khurana SM (2003) Electronic microscopic studies of a Potyvirus infecting Datura stramonium. XXVI annual conference on electron microscopy and allied fields, 16–18th Apr 2003, CPRI, Shimla (India). pp 7–8 Jackson AO, Goodin MM, Ganesan U, Bragg J (2011) Nucleorhabdovirus. Rhabdoviridae. In: The Springer index of viruses. Springer, New York, pp 1741–1745. https://doi.org/10.1007/978-0-387-95919-1_281 Jorda C, Font I, Martinez P, Juarez M, Ortega A, Lacasa A (2001) Current status and new natural hosts of Tomato yellow leaf curl virus (TYLCV) in Spain. Plant Dis 85:445 Kahn RP, Bartels R (1968) The Colombian datura virus – a new virus in the Potato virus Y group. Phytopathology 58:587–592 Kaur C, Kumar S, Raj SK (2015) Association of a distinct strain of Chilli veinal mottle virus with mottling and distortion disease of Datura inoxia in India. Arch Phytopathol Plant Protect 48:545–554 Mackie AE, Rodoni BC, Barbetti MJ, McKirdy SJ, Jones RAC (2016) Potato spindle tuber viroid: alternative host reservoirs and strain found in a remote subtropical irrigation area. Eur J Plant Pathol 145:433–446 Matousek J, Piernikarczyk RJJ, Dedic P, Mertelík J, Uhlířová K, Duraisamy G, Orctová L, Kloudová K, Ptáček J, Steger G (2014) Characterization of Potato spindle tuber viroid (PSTVd) incidence and new variants from ornamentals. Eur J Plant Pathol 138:93–101 Mohammed HS, El Siddig MA, El Hussein AA, Navas-Castillo J, Fiallo-Olive E (2017) First report of Datura innoxia as a natural host of Watermelon chlorotic stunt virus in Sudan. Plant Dis 101:1334 Mohammed HS, El Siddig MA, El Hussein AA, Navas-Castillo J, Fiallo-Olivé E (2018) Complete genome sequence of datura leaf curl virus, a novel begomovirus infecting Datura innoxia in Sudan, related to begomoviruses causing tomato yellow leaf curl disease. Arch Virol 163(1):273–275 Moodley V, Gubba A, Mafongoya PL (2016) Occurrence of Tomato chlorosis virus (ToCV) on Datura stramonium near tomato crops (Solanum lycopersicum) in South Africa. Plant Dis 100:1512 Papayiannis L, Katis NI, Idris AM, Brown JK (2011) Identification of weed hosts of Tomato yellow leaf curl virus in Cyprus. Plant Dis 95:120–125 Prasada Rao RDVJ, Yaraguntaiah RC (1976) Natural occurrence of Potato virus Y on Datura metel. Curr Sci 45:467 Raj SK, Srivastava A, Chandra G, Singh BP (1999) Natural occurrence of cucumber mosaic cucumovirus in Datura innoxia in India. EPPO Bull 29:455–457 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Roggero P, Accotto GP, Ciuffo M, Lenzi R, Desbiez C, Lecoq H, Bosco D, Huang X, Gu Q (2000) First report of Tobacco vein banding mosaic virus in China (Xian, Shaanxi Province) in Datura stramonium and tobacco. Plant Dis 84:1152.4 Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct Tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207 Taiwo MA, Hughes J’dA, Akomah AR (2004) Natural infection of D. stramonium L. by an unusual strain of PVMV in Nigeria. Phytopathology 94:S101 Taiwo MA, Hughes JA, Akomah AR (2006) Natural infection of Datura stramonium L. by an unusual strain of Pepper veinal mottle virus Genus Potyvirus in Nigeria. Plant Pathol 5:401–404 Thomas JE, Behncken GM (1983) A Rhabdovirus infecting tomato and Datura stramonium in south-east Queensland. Abstr. 4th Int. Congr. Pl. Path., Melbourne, p 123 Thomas JE, Dietzgen RG (1991) Characterisation of Datura yellow vein virus, a newly described rhabdovirus from Australia. Ann Appl Biol 118:339–349 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Rhabdoviridae. J Gen Virol 99:447–448 Webster CG, Jensen CE De Rivera-Vargas LI, Rodrigues JCV, Mercado W, Frantz G, Mellinger HC, Adkins S (2013) First report of Tomato chlorotic spot virus (TCSV) in tomato, pepper, and jimsonweed in Puerto Rico. Plant Health Progress No. August pp. PHP-2013-0812-01-BR Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

Daucus carota (Carrot)

841

Daucus carota (Carrot) Family: Apiaceae

Vegetable

Ageratum enation virus Taxonomic position Genus: Begomovirus

(AEV)

D Family: Geminiviridae

AEV infection in plants of Daucus carota was reported from India (Kumar et al. 2013). The virusinfected carrot plants exhibit symptoms of yellow mosaic and shortening of leaf petiole. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of AEV, refer to Ageratum spp.

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

(AMV)

Family: Bromoviridae

AMV infection in plants of Daucus carota was reported from India, southern Washington (USA), and Germany (Campbell and Melugin 1971; Wolf and Schmelzer 1973; Douine 1976; Howell and Mink 1981; Rafiq et al. 2008). The virus-infected carrot plants exhibit symptoms of spotting, mottling, mosaic, and dwarfing. The virus is transmitted by several species of aphids in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Broad bean wilt virus Taxonomic position Genus: Fabavirus

(BBWV)

Family: Secoviridae

BBWV (but without specifying either of the currently recognized species BBWV-1 or BBWV-2) infection in plants of Daucus carota was reported from Czechoslovakia (Chod and Jokes 1986). The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BBWV, refer to Vicia faba.

Carrot Ch virus 1 Taxonomic position Genus: Chordovirus

(CChV–1)

Family: Betaflexiviridae

Geographical distribution CChV-1 infection in plants of Daucus carota was reported from the UK (Adams et al. 2014).

842

Daucus carota (Carrot)

Symptoms and host(s) The virus-infected carrot plants exhibit root necrosis symptoms. Transmission The virus is mechanically sap-transmissible. Virion properties and genome The virions are flexuous filaments, usually 12–13 nm in diameter (range 10–15 nm) and from 600 to over 1000 nm in length. The genome consists of a single molecule of linear single-stranded RNA of 8155 nt (KF533711) (Adams et al. 2014).

Carrot Ch virus 2 Taxonomic position Genus: Chordovirus

(CChV–2)

Family: Betaflexiviridae

Geographical distribution CChV-2 infection in plants of Daucus carota was reported from the UK (Adams et al. 2014). Symptoms and host(s) The virus-infected carrot plants exhibit root necrosis symptoms. Transmission The virus is mechanically sap-transmissible. Virion properties and genome The virions are flexuous filaments, usually 12–13 nm in diameter (range 10–15 nm) and from 600 to over 1000 nm in length. The genome consists of a single molecule of linear single-stranded RNA of 8047 nt (KF533710) (Adams et al. 2014).

Carrot cryptic virus Taxonomic position Genus: Alphapartitivirus

(CaCV)

Family: Partitiviridae

Geographical distribution CaCV infection in plants of Daucus carota was reported from Germany, but has similarities to Carrot temperate virus 4, from Japan (Willenborg et al. 2009). Symptoms and host(s) The virus-infected carrot plants do not exhibit any symptoms. Transmission The virus is not transmitted by an insect vector. The virus is not mechanically sap-transmissible and no contact transmission. The virus is not graft-transmissible. The virus is transmitted by pollen to the seed.

Daucus carota (Carrot)

843

Virion properties and genome The virions are non-enveloped isometric and 30 nm in diameter. The genome consists of two segments of double-stranded RNA. RNA1 consists of 1971 bp (FJ550604) and RNA2 of 1776 bp (FJ550605) (Willenborg et al. 2009; Vainio et al. 2018).

Carrot latent virus

(CtLV)

Taxonomic position CtLV is a tentative member of the family Rhabdoviridae Geographical distribution CtLV infection was first reported in plants of Daucus carota from Japan by Ohki et al. (1978). The virus spreads in Japan, Europe, and the UK. Symptoms and host(s) The virus-infected carrot plants exhibit symptomless infections. Transmission The virus is transmitted by the aphid vectors, Brachycaudus heraclei and Semiaphis heraclei. The virus is not transmissible by mechanical inoculation. Virion properties and genome The virions are rhabdo- or bullet-shaped and enveloped, with a clear modal length of 220 nm and 70 nm wide. No sequence is currently available.

Carrot mottle mimic virus Taxonomic position Genus: Umbravirus

(CMoMV)

Family: Tombusviridae

Geographical distribution CMoMV infection was first reported in plants of Daucus carota from New South Wales, Australia, by Gibbs (1991, 1995). The virus spreads in Australia, New Zealand, and the USA (Gibbs et al. 1996b; Tang et al. 2009). Symptoms and host(s) The virus-infected carrot plants exhibit systemic yellowing, mottling, and stunting symptoms. Transmission The virus is transmitted by an aphid vector probably Cavariella aegopodii in a circulative, nonpropagative manner. CMoMV requires, for vector transmission, a helper virus (Carrot red leaf virus, CtRLV, a Polerovirus). The virus is transmissible by mechanical sap-inoculation to less than three families. Experimentally infected plants mostly show vein necrosis.

D

844

Daucus carota (Carrot)

Virion properties and genome CMoMV has no conventional virus particles and they lack a coat protein ORF but is encapsidated within particles of a helper polerovirus (CtRLV). The genome is positive-sense single-stranded RNA of 4201 nt (U57305 = NC_001726) (Gibbs et al. 1996b).

Carrot mottle virus Taxonomic position Genus: Umbravirus

(CMoV)

Family: Tombusviridae

Geographical distribution CMoV infection in plants of Daucus carota was originally described from Great Britain (Watson et al. 1964). The virus occurs worldwide, including Europe, New Zealand, Japan, Israel, North America, the UK, Mauritius, Germany, and Australia (Wolf and Schmelzer 1973; Murant et al. 1985; Menzel et al. 2008; Gungoosingh-Bunwaree et al. 2009). Symptoms and host(s) The virus-infected carrot plants exhibit yellowing or reddening and plants are also stunted. The natural host range of this virus is restricted to the Amaranthaceae, Chenopodiaceae, Solanaceae, and Umbelliferae families. Transmission The virus is transmitted through aphid vectors, Cavariella aegopodii and Myzus persicae, in a circulative non-propagative manner (Murant et al. 1969; Waterhouse and Murant 1983; Naseem et al. 2016). The virus is retained when the vector moults and does not multiply in the vector, and is not vertically transmitted to progeny. The virus is mechanically sap-transmissible, but not transmitted through seed or pollen. Virion properties and genome The viral genome lacks a coat protein ORF and conventional virions are not produced but the genome is encapsidated by a helper virus (Carrot red leaf virus, a Polerovirus). The genome is a positive-sense ssRNA of 4193 nt (FJ188473 = NC_011515) which contains four open reading frames (ORF). The ORF-1 encodes a 34.2 kDa protein. A signal sequence, GGATTTT (starting at 980 nucleotide position), is present in CMoV for ribosome frameshift from ORF-1 to ORF-2, resulting in production of a 95.9 kDa protein. The ORF-3 and ORF-4 overlap each other, and they are separated by 174 nt intergenic region, which may contain promoter sequences for the subgenomic transcript used to translate the ORF-3 and ORF-4 as 28.9 kDa and 30.8 kDa protein, respectively (Murant et al. 1985; Gibbs et al. 1996a; Menzel et al. 2008).

Carrot necrotic dieback virus Taxonomic position Genus: Sequivirus

(CNDV)

Family: Secoviridae

Daucus carota (Carrot)

845

Geographical distribution CNDV infection in plants of Daucus carota was reported from Germany (Menzel and Vetten 2008a). Symptoms and host(s) The virus-infected carrot plants exhibit necrotic dieback symptoms. Transmission The virus is transmitted by aphid vectors in a semi-persistent manner, however, it is dependent on the presence of a helper virus in the genus Waikavirus. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome consists of a single molecule of linear positive-sense, single-stranded RNA of 9905 nt (EU980442 = NC_038320) that encodes a single large polyprotein (Sanfacon et al. 2009; Reavy 2011; Sanfacon 2015; Thompson et al. 2017).

Carrot red leaf virus

(CtRLV)

Synonyms Anthricus strain; Parsnip yellow fleck virus Taxonomic position Genus: Polerovirus

Family: Luteoviridae

Geographical distribution CtRLV infection in plants of Daucus carota was first reported from Great Britain (Watson et al. 1964). The virus occurs worldwide, including Mauritius, Australia, Canada, Germany, Japan, New Zealand, the UK, and the USA (Ohki et al. 1979; Waterhouse and Murant 1981; Waterhouse 1985; Marco 1993; Morton et al. 2003; Huang et al. 2005; Gungoosingh-Bunwaree et al. 2009; Tang et al. 2009). Symptoms and host(s) The virus-infected carrot plants exhibit reddening and yellowing of leaves and some stunting of the plants. The natural host range of this virus is cultivated in wild carrot, parsley (Petroselinum crispum), some cow parsley (Anthriscus sylvestris), and hogweed (Heracleum sphondylium). Transmission The virus is not mechanically sap-transmissible but is transmitted by the aphids Caraviella aegopodii, Myzus persicae, and Semiaphis heraclei in a circulative, non-propagative manner (Naseem et al. 2016). The virus is retained when the vector moults and does not multiply in the vector and is not transmitted congenitally to the progeny of the vector. The virus is not seed-transmitted (Waterhouse and Murant 1983). Virion properties and genome The virions are 25–30 nm in diameter and hexagonal in outline and have no envelope. The genome is a positive-sense ssRNA of 5723 nt (NC_006265, AY695933). The 30 terminus has neither a poly(A) tract nor a tRNA-like structure. The 50 terminus has a genome-linked protein (VPg) (of about 17 kDa). The

D

846

Daucus carota (Carrot)

genome consists of six open reading frames (ORFs) (Murant et al. 1985; Morton et al. 2003; Huang et al. 2005).

Carrot temperate virus 1 Taxonomic position Genus: Unassigned

(CteV-1)

Family: Partitiviridae

Geographical distribution CteV-1 infection was first reported in plants of Daucus carota from Japan by Natsuaki et al. (1983). The virus spreads in Japan. Symptoms and host(s) The virus-infected carrot plants exhibit symptomless infections. Transmission The virus is not transmissible by mechanical sap-inoculation, and also not by grafting. Virion properties and genome The virions are isometric, non-enveloped, and 30 nm in diameter. The genome typically consists of two dsRNA segments (Vainio et al. 2018).

Carrot temperate virus 2 Taxonomic position Genus: Unassigned

(CteV-2)

Family: Partitiviridae

Geographical distribution CteV-2 infection was first reported in plants of Daucus carota from Japan (Natsuaki et al. 1990). The virus spreads in Japan. Symptoms and host(s) The virus-infected carrot plants exhibit symptomless infections. Transmission The virus is not transmissible by mechanical sap-inoculation, and not transmissible by grafting. Virion properties and genome The virions are isometric, non-enveloped, and 30 nm in diameter. The genome typically consists of two dsRNA segments (Vainio et al. 2018).

Carrot temperate virus 3 Taxonomic position Genus: Unassigned

(CteV-3)

Family: Partitiviridae

Daucus carota (Carrot)

847

Geographical distribution CteV-3 infection was first reported in plants of Daucus carota from Japan (Natsuaki et al. 1990). The virus spreads in Japan. Symptoms and host(s) The virus-infected carrot plants exhibit symptomless infections. Transmission The virus is not transmissible by mechanical sap-inoculation, and not transmissible by grafting. Virion properties and genome The virions are isometric, non-enveloped, and 30 nm in diameter. The genome typically consists of two dsRNA segments (Vainio et al. 2018).

Carrot temperate virus 4 Taxonomic position Genus: Unassigned

(CteV-4)

Family: Partitiviridae

Geographical distribution CteV-3 infection was first reported in plants of Daucus carota from Japan (Natsuaki et al. 1990). The virus spreads in Japan. Symptoms and host(s) The virus-infected carrot plants exhibit symptomless infections. Transmission The virus is not transmissible by mechanical sap-inoculation, and not transmissible by grafting. Virion properties and genome The virions are isometric, non-enveloped, and 30 nm in diameter. The genome typically consists of two dsRNA segments (Vainio et al. 2018).

Carrot thin leaf virus Taxonomic position Genus: Potyvirus

(CTLV)

Family: Potyviridae

Geographical distribution CTLV infection in plants of Daucus carota was reported from California, Idaho, and Washington (USA) (Howell and Mink 1976a, 1981; Falk et al. 1991). Symptoms and host(s) The virus-infected carrot plants show leaflets that are thinner than normal, giving the plant an overall unusual appearance. In general, leaflets are threadlike and twisted, giving the foliage a narrow and distorted appearance. Leaves also may exhibit faint mottling and yellow vein-banding. When plants are

D

848

Daucus carota (Carrot)

infected at a young age, the leaflets may be extremely thin, hence the name of disease. Symptoms vary by carrot cultivar and the growth stage when infection occurred (Howell and Mink 1976b). The natural host range of this virus is cultivated in wild carrot and the weed hemlock (Conium maculatum). Transmission The virus is transmitted by aphids Cavariella aegopodii and Myzus persicae in a non-persistent manner, and is sap-transmissible to some plant species belonging to the families Apiaceae, Chenopodiaceae, Asteraceae, Fabaceae, and Solanaceae (Xu et al. 2014). Virion properties and genome The virions are non-enveloped, flexuous, filaments particles measuring 736 nm in length and 11 nm in width. The genome consists of a single molecule of linear, positive-sense ssRNA of 9491 nt (JX156434 = NC_025254) (Howell and Mink 1976b; Xu et al. 2014; Revers and Garcia 2015; Wylie et al. 2017).

Carrot torradovirus 1 Taxonomic position Genus: Torradovirus

(CaTV1)

Family: Secoviridae

Geographical distribution CaTV1 infection in plants of Daucus carota was reported from the UK (Adams et al. 2014; Fox et al. 2017; Rozado-Aguirre et al. 2016, 2017a, b). Symptoms and host(s) The virus-infected carrot plants exhibit necrotic symptoms. Transmission The virus is demonstrated to be transmitted by at least three different species of aphids (Rozado-Aguirre et al. 2016). The virus is also transmissible by mechanical sap-inoculation. Virion properties and genome The virions are isometric, non-enveloped of two types but similar in size 25–30 nm in diameter, and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is a bipartite, linear positive-sense, single-stranded RNA. RNA1 is 6944 nt (KF533719 = NC_025479) and RNA2 is 4995 nts (KF533720 = NC_025480) (Sanfacon et al. 2009; Adams et al. 2014; Sanfacon 2015; van der Vlugt et al. 2015; Rozado-Aguirre et al. 2017a, b; Thompson et al. 2017).

Carrot virus S

(CarVS)

Taxonomic position CarVS is a tentative member of the genus Carlavirus and family Betaflexiviridae Geographical distribution CarVS infection in plants of Daucus carota was reported from Germany (Menzel and Vetten 2008b).

Daucus carota (Carrot)

849

Transmission The virus is mechanically sap-transmissible. Virion properties and genome A partial genome sequence of 2235 nt from the 30 end of the genome is available (EU881919) (Menzel and Vetten 2008b); the full genome is expected to be c.8.5–9 kb.

D

Carrot virus Y

(CarVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution CarVY infection in plants of Daucus carota was reported from Australia (Latham and Jones 2002; Moran et al. 2002; Latham et al. 2004). Symptoms and host(s) This virus infects carrots at any stage. The typical symptoms include chlorotic mottling, marginal necrosis or reddening and generalized chlorosis of leaves. The most severe symptom is the increased subdivision of leaflets giving a feathery appearance, and plants are also stunted. The virus causes severe root symptoms in carrots including shortened roots, knobbliness, and severe distortion, particularly if plants are infected at an early growth stage. Later infection (carrot plants infected more than 6 weeks after germination) produces milder symptoms of thinner carrots with only slight distortion but still with a substantial overall yield loss (Latham and Jones 2004). Transmission The virus spreads by aphids, Myzus persicae and other aphids in a non-persistent manner; M. persicae is most efficient. The virus also infects several species in the Apiaceae by sap-inoculation. Seedtransmission is unknown (Jones et al. 2005, 2006). Virion properties and genome The virions are non-enveloped, flexuous, filaments about 11 nm in width and 770 nm in length. The genome consists of a single molecule of linear positive-sense ssRNA, of c.9.5–10 kb. The partial sequence of 1389 nt from the NIa/NIb region (DQ174243) and 1754 nt of the partial NIb polymerase and complete coat protein (AF203537, AF203538, and AF203539) are available (Moran et al. 2002; Revers and Garcia 2015; Wylie et al. 2017).

Carrot yellow leaf virus Taxonomic position Genus: Closterovirus

(CYLV)

Family: Closteroviridae

Geographical distribution CYLV was first reported in plants of Daucus carota from Japan by Yamashita et al. (1976). The virus spreads in Japan, the Netherlands, and the UK (Van Dijk and Bos 1996; Adams et al. 2014).

850

Daucus carota (Carrot)

Symptoms and host(s) The virus-infected carrot plants exhibit necrosis, leaf yellowing, and reddening symptoms. A wild plant, hogweed (Heracleum sphondylium), is a natural host of CYLV (Adams et al. 2014). Transmission The virus is transmitted by the aphid vectors, Cavariella aegopodii, C. pastinacae, and C. theobaldi, in a semi-persistent manner. CYLV can help the vector transmission of another virus (Heracleum latent virus, a Vitivirus). The virus is transmissible by mechanical sap-inoculation (with difficulty). Experimentally infected plants mostly show yellowing and necrosis symptoms. Virion properties and genome The virions are flexuous filaments, with a clear modal length of 1650 nm and 12 nm wide. The genome consists of a single molecule of linear positive-sense ssRNA of 16,354 nt (FJ869862). The genome contains ten open reading frames (ORFs), and their organization is similar to Beet yellow stunt virus; however, ORF-2 and ORF-3 are in a reversed order (Menzel et al. 2009; Agranovsky and Lesemann 2011).

Celery mosaic virus

(CeMV)

Synonyms Western celery mosaic virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

CeMV infection in plants of Daucus carota was reported from the USA, Australia, Japan, and Germany (Iwaki and Komuro 1970; Wolf and Schmelzer 1973; Howell and Mink 1981; Chod 1984; Traicevski et al. 1999). The virus-infected carrot plants exhibit spotting, mottling, mosaic, malformation, and dwarfing symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CeMV, refer to Apium graveolens.

Citrus exocortis viroid Taxonomic position Genus: Pospiviroid

(CEVd)

Family: Pospiviroidae

CEVd infection in plants of Daucus carota was reported from Spain (Fagoaga and Duran-Vila 1996). The viroid is mechanically sap-transmissible and is also transmissible through contaminated tools. For more details of CEVd, refer to Citrus spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

Daucus carota (Carrot)

851

The subgroup II CMV was identified causing a chlorotic mottle disease of Daucus carota in northern Uttar Pradesh, India, and Japan (Iwaki and Komuro 1970; Wolf and Schmelzer 1973; Afreen et al. 2009). The virus-infected carrot plants exhibit chlorotic mottling, mosaic, and yellowing symptoms. The virus is transmitted by a number of aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of host plants. For more details of CMV, refer to Cucumis sativus.

D Lettuce infectious yellows virus Taxonomic position Genus: Crinivirus

(LIYV)

Family: Closteroviridae

LIYV infection in plants of Daucus carota was reported from the Southwestern USA (Duffus et al. 1986). The virus-infected carrot plants exhibit yellowing symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a semi-persistent manner. The virus is not mechanically saptransmissible. For more details of LIYV, refer to Lactuca sativa.

Parsnip yellow fleck virus Taxonomic position Genus: Sequivirus

(PYFV)

Family: Secoviridae

PYFV infection in plants of Daucus carota was reported from the UK and the Netherlands (Van Dijk and Bos 1985; Morgan 2004). The virus-infected carrot plants initially exhibit the first symptoms in late May and early June resulting in severely stunted plants and the death of many individual plants. Later in the season, larger plants develop mottled foliage that is discolored with yellow flecks. Plants infected with virus may develop secondary and/or misshapen roots, and throughout the season, the tops of infected plants can develop dieback and rot. The virus is transmitted by the willow-carrot aphid, Cavariella aegopodii, in a semi-persistent manner, however, it is dependent on the presence of a helper virus in the genus Waikavirus. The virus is also transmissible by mechanical sap-inoculation, but is not transmitted by seed and pollen. For more details of PYFV, refer to Pastinaca sativa.

Pedilanthus leaf curl virus Taxonomic position Genus: Begomovirus

(PeLCV)

Family: Geminiviridae

PeLCV infection in plants of Daucus carota was reported from India (Saritha et al. 2017). The virusinfected carrot plants exhibit symptoms of downward curling without yellowing and reduced vigor. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of PeLCV, refer to Pedilanthus tithymaloides.

852

Daucus carota (Carrot)

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Daucus carota was reported from India (Sivalingam et al. 2011). The virus-infected carrot plants exhibit yellow vein symptoms starting from the leaf margins which gradually intensify until the entire lamina appears yellow within a week, and severely affected plants showed reduced lamina size and mild curling. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner, and also by grafting. For more details of ToLCNDV, refer to Solanum lycopersicum.

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV infection in plants of Daucus carota was reported from Australia (Parry and Persley 2005). The virus-infected carrot plants do not exhibit any symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation, and its host range was restricted mostly to Cucurbitaceae, Leguminosae, and Chenopodiaceae. For more details of WMV, refer to Citrullus lanatus.

References Adams IP, Skelton A, Macarthur R, Hodges T, Hinds H, Flint L, Nath PD, Boonham N, Fox A (2014) Carrot yellow leaf virus is associated with carrot internal necrosis. PLoS One 9(11):E109125 Afreen B, Khan AA, Naqvi QA, Kumar S, Pratap D, Snehi SK, Raj SK (2009) Molecular identification of a Cucumber mosaic virus subgroup II isolate from carrot (Daucus carota) based on RNA3 genome sequence analyses. J Plant Dis Protect 116(5):193–199 Agranovsky AA, Lesemann D-E (2011) Closterovirus. Closteroviridae. In: The Springer index of viruses. Springer, New York, pp 327–333. https://doi.org/10.1007/978-0-387-95919-1_50 Campbell RN, Melugin SA (1971) Alfalfa mosaic virus strains from carrot and parsley. Plant Dis Reptr 55:322–325 Chod J (1984) Detection of celery mosaic virus in carrot variety Nantes. Sbbornik – Uvitz Ochrana Rostlin 20:91–96 Chod J, Jokes M (1986) Incidence of broad bean wilt virus on carrot (Daucus carota) in Czechoslovakia. Sbornik UVTIZ, Ochrana Rostlin 22(4):241–248 Douine LA (1976) New strain of Lucerne mosaic virus isolated from carrot (Daucus carota L.) in southeastern France. Ann de Phytopath 89:343–346 Duffus JE, Larsen RC, Liu HY (1986) Lettuce infectious yellows virus – a new whitefly transmitted virus. Phytopathology 76:97–100 Fagoaga C, Duran-Vila N (1996) Naturally occurring variants of citrus exocortis viroid in vegetable crops. Plant Pathol 45:45–53 Falk BW, Davis RM, Piechocki M (1991) Identification of Carrot thin leaf virus in California carrots. Plant Dis 75:319 Fox A, Rozado Z, Adams IP, Skelton A, Dickinson M, Boonham N (2017) Investigating the viral causes of carrot internal necrosis. Acta Hortic 1153:245–250 Gibbs MJ (1991) Grad. Dip. Thesis, Aust. NAT Univ Gibbs MJ (1995) The genome of carrot mottle mimic umbravirus and the evolution of the carmo and sobemo virus families. D Phil thesis, University of Oxford, Oxford Gibbs MJ, Cooper JI, Waterhouse PM (1996a) The genome organisation and affinities of an Australian isolate of carrot mottle umbravirus. Virology 224:310–313

Daucus carota (Carrot)

853

Gibbs MJ, Ziegler A, Robinson DJ, Waterhouse PM, Cooper JI (1996b) Carrot mottle mimic virus (CMoMV), a second umbravirus associated with carrot motley dwarf disease recognized by nucleic acid hybridization. Mol Plant Pathol. On-line URL http://www.bspp.org.uk/mppol/1996/1111gibbs Gungoosingh-Bunwaree A, Menzel W, Winter S, Vally V, Seewoogoolam R, BeniMadhu SP, Vetten HJ (2009) First report of Carrot red leaf virus and Carrot mottle virus, causal agents of carrot motley dwarf, in carrot in Mauritius. Plant Dis 93(11):1218 Howell WE, Mink GI (1976a) Carrot thin leaf potyvirus. First reported in Daucus carota; from Washington State, USA. Phytopathology 66:949 Howell WE, Mink GI (1976b) Host range, purification and properties of a flexuous rod shaped virus isolated from carrot. Phytopathology 66:949–953 Howell WE, Mink GI (1981) Viruses isolated from wild carrot and poison hemlock. Plant Dis 65:277–279 Huang LE, Naylor M, Pallet DW, Reeves J, Cooper JI, Wang H (2005) The complete genome sequence, organization and affinities of Carrot red leaf virus. Arch Virol 150:1845–1855 Iwaki M, Komuro Y (1970) Viruses isolated from carrot. 1. Celery mosaic virus and Cucumber mosaic virus. Ann Phytopathol Soc Jpn 36:36–42 Jones RAC, Smith LJ, Gajda BE, Smith TN, Latham LJ (2005) Further studies on Carrot virus Y: host range, symptomatology, search for resistance and tests for seed transmissibility. Aust J Agric Res 56:859–868 Jones RAC, Smith LJ, Smith TN, Latham LJ (2006) Relative abilities of different aphid species to act as vectors of Carrot virus Y. Australian Plant Pathology 35:23–27 Kumar J, Gunapati S, Singh SP, Gadre R, Sharma NC, Tuli R (2013) Molecular characterization and pathogenicity of a carrot (Daucus carota) infecting begomovirus and associated betasatellite from India. Virus Res 178:478–485 Latham LJ, Jones RAC (2002) Carrot virus Y. In: Davis M, Raid R (eds) Compendium of umbelliferous diseases. American Phytopathological Society Press, Minnesota, p 53 Latham LJ, Jones RAC (2004) Carrot virus Y: symptoms, losses, incidence, epidemiology and control. Virus Res 100:89–99 Latham LJ, Traicevski V, Persley DM, Wilson CR, Tesoriero L, Coles R, Jones RAC (2004) Distribution and incidence of Carrot virus Y in Australia. Australas Plant Pathol 33:83–86 Marco S (1993) Carrot red leaf virus in Israel. Phytoparasitica 21:123 Menzel W, Vetten HJ (2008a) Complete nucleotide sequence of an isolate of the Anthriscus strain of Parsnip yellow fleck virus. Arch Virol 153(11):2173–2175 Menzel W, Vetten HJ (2008b) Partial nucleotide sequence of a carlavirus from carrot (Daucus carota L.). Virus Genes 37:432–433 Menzel W, Maiss E, Vetten HJ (2008) Complete nucleotide sequence of a carrot isolate of Carrot mottle virus from Germany. Arch Virol 153:2163–2165 Menzel W, Goetz R, Lesemann DE, Vetten HJ (2009) Molecular characterization of a closterovirus from carrot and its identification as a German isolate of Carrot yellow leaf virus. Arch Virol 154:1343–1347 Moran J, van Rijswijk B, Traicevski V, Kitajima EW, Mackenzie AM, Gibbs AJ (2002) Potyviruses, novel and known, in cultivated and wild species of the family Apiaceae in Australia. Arch Virol 147:1855–1867 Morgan D (2004) Parsnip yellow fleck virus: development of a disease management strategy. Horticultural Development Company. Available: http://www.hdc.org.uk/project/parsnip-yellow-fleck-virus-development-disease-managementstrategy-4. Accessed 25 June 2014 Morton A, Spence NJ, Boonham N, Barbara DJ (2003) Carrot red leaf associated RNA in carrots in the United Kingdom. Plant Pathol 52:795 Murant AF, Goold RA, Roberts IM, Cathro J (1969) Carrot mottle – a persistent aphid-borne virus with unusaual properties and particles. J Gen Virol 4:329–341 Murant AF, Waterhouse PM, Raschke JH, Robinson DJ (1985) Carrot red leaf and Carrot mottle virus: observations on the composition of the particles in single and mixed infections. J Gen Virol 66:1575–1579 Naseem MT, Ashfaq M, Khan AM, Kiss Z, Akhtar KP, Mansoor S (2016) Transmission of viruses associated with carrot motley dwarf by Myzus persicae. J Plant Pathol 98:581–585 Natsuaki T, Yamashita S, Doi Y, Okuda S, Teranaka M (1983) Radish yellow edge virus, a seed-borne virus with doublestranded RNA, of a possible new group. Ann Phytopathol Soc Jpn 49:593–599 Natsuaki T, Muroi Y, Okuda S, Teranaka M (1990) Ann Phytopath Soc Japan 56:354 Ohki ST, Doi Y, Yora K (1978) Carrot latent virus: a new rhabdovirus of carrot. Ann Phytopath Soc Japan 44:202–204 Ohki ST, Doi Y, Yora K (1979) Small spherical virus particles found in carrot plants infected with Carrot red leaf virus. Ann Phytopathol Soc Jpn 45:74–76 Parry JN, Persley DM (2005) Carrot as a natural host of Watermelon mosaic virus. Australian Plant Pathology 34:283–284 Rafiq A, Ali S, Jahan T, Naqvi QA (2008) Virus causing yellow net disease on carrot (Daucus carota L.) identified as Alfalfa mosaic virus. National Academy Science Letters 31(1/2):39–43 Reavy B (2011) Sequivirus. Sequiviridae. In: The Springer index of viruses. Springer, New York, pp 1771–1774. https:// doi.org/10.1007/978-0-387-95919-1_288 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Rozado-Aguirre Z, Adams I, Collins L, Fox A, Dickinson M, Boonham N (2016) Detection and transmission of Carrot torrado virus, a novel putative member of the Torradovirus genus. J Virol Methods 235:119–124

D

854

Deinbollia borbonica

Rozado-Aguirre Z, Marais A, Svanella-Dumas L, Faure C, Latour F, Villeneuve F, Dickinson M, Fox A, Boonham N, Candresse T (2017a) First report of Carrot torradovirus 1 (CaTV1) a member of the Torradovirus genus, infecting carrots in France. Plant Dis 101:1333 Rozado-Aguirre Z, Adams I, Fox A, Dickinson M, Boonham N (2017b) Complete sequence and genomic annotation of Carrot torradovirus 1. Arch Virol 162:2815–2819 Sanfacon H (2015) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. In: eLS. Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000764.pub3. http://www.els.net Sanfacon H, Wellink J, Le Gall O, Karasev A, van der Vlugt R, Wetzel T (2009) Secoviridae: a proposed family of plant viruses within the order Picornavirales that combines the families Sequiviridae and Comoviridae, the unassigned genera Cheravirus and Sadwavirus, and the proposed genus Torradovirus. Arch Virol 154(5):899–907 Saritha RK, Shrawan S, Kalia P, Jain RK, Baranwal VK (2017) Association of Pedilanthus leaf curl virus and satellites with carrot (Daucus carota) in India. Plant Dis 101:843 Sivalingam PN, Sumiya KV, Malathi VG (2011) Carrot as a new host for a begomovirus: yellow mosaic disease of carrot reported in India. New Disease Reports 23:34 Tang J, Quinn BD, Clover GRG (2009) First report of Carrot red leaf virus-associated RNA co-infecting carrot with Carrot red leaf virus and carrot mottle mimic virus to cause carrot motley dwarf disease in New Zealand. Aust Plant Dis Notes 4:15–16 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Secoviridae. J Gen Virol 98:529–531 Traicevski V, Schreurs T, Rodoni B, Moran J (1999) Celery mosaic potyvirus occurring naturally in cultivated Apiaceae in Victoria, Australia. Australas Plant Pathol 28:92 Vainio EJ, Chiba S, Ghabrial SA, Maiss E, Roossinck M, Sabanadzovic S, Suzuki N, Xie J, Nibert M, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Partitiviridae. J Gen Virol 99:17–18 van der RAA V, Verbeek M, Dullemans AM, Wintermantel WM, Cuellar WJ, Fox A, Thompson JR (2015) Torradoviruses. Annu Rev Phytopathol 53:485–512 Van Dijk P, Bos L (1985) Viral dieback of carrot and other umbelliferae caused by the Anthriscus strain of parsnip yellow fleck virus, and its distinction from carrot motley dwarf. Neth J Plant Pathol 91:169–187 Van Dijk P, Bos L (1996) Carrot yellow leaf virus. In: Brunt AA, Crabtree K, Dallwitz MJ, Gibbs AJ, Watson L (eds) Viruses of plants. Descriptions and lists from the VIDE database. CAB International, pp 325–326 Waterhouse PM (1985) Isolation and identification of Carrot red leaf virus from carrot and dill growing in the Australian Capital Territory. Australas Plant Pathol 14:32–34 Waterhouse PM, Murant AF (1981) Purification of Carrot red leaf virus and evidence from four serological tests for its relationship to luteoviruses. Ann Appl Biol 97:91–204 Waterhouse PM, Murant AF (1983) Further evidence one the nature of the dependence of Carrot mottle virus on Carrot red leaf virus for transmission by aphids. Ann Appl Biol 103:455–464 Watson MA, Serjeant EP, Lennon EA (1964) Carrot motley dwarf and parsnip mottle viruses. Ann Appl Biol 54:163–166 Willenborg J, Menzel W, Vetten H-J, Maiss E (2009) Molecular characterization of two alphacryptovirus dsRNAs isolated from Daucus carota. Arch Virol 154(3):541–543 Wolf P, Schmelzer K (1973) Virus diseases of carrot (Daucus carota L.). Acta Phytopathol Acad Sci Hung 8:311–327 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354 Xu D, Liu H-Y, Li F, Tian T, Li R (2014) Characterizations of Carrot thin leaf virus based on host reactions and complete genomic sequences. Eur J Plant Pathol 138:15–22 Yamashita S, Ohki S, Doi Y, Yora K (1976) Identification of two viruses associated with the carrot yellow leaf syndrome. Ann Phytopathol Soc Jpn 42:382–383

Deinbollia borbonica Family: Sapindaceae

Deinbollia mosaic virus Taxonomic position Genus: Begomovirus

Tree for wood

(DbMV)

Family: Geminiviridae

Delphinium spp. (Larkspur)

855

Geographical distribution DbMV infection in plants of Deinbollia borbonica was reported from Kenya and Tanzania (Kyallo et al. 2017a). Symptoms and host(s) The virus-infected Deinbollia plants exhibit yellow mosaic symptoms. The virus infects a restricted host range of Solanaceae and Euphorbiaceae species (Kyallo et al. 2017b). Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm, with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2759 nt (KT878829 = NC_029804) and DNA-B of 2702 nt (KT878828 = NC_029803) (Briddon 2001; Brown et al. 2015; Kyallo et al. 2017a; Zerbini et al. 2017).

References Briddon RW (2001) Begomovirus. Geminiviridae. The Springer index of plant viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Kyallo M, Sseruwagi P, Skilton RA, Ochwo-Ssemakula M, Wasswa P, Ndunguru J (2017a) Deinbollia mosaic virus: a novel Begomovirus infecting the sapindaceous weed Deinbollia borbonica in Kenya and Tanzania. Arch Virol 162:1393–1396 Kyallo M, Ateka EM, Sseruwagi P, Ascencio-Ibáñez JT, Ssemakula MO, Skilton R, Ndunguru J (2017b) Infectivity of Deinbollia mosaic virus, a novel weed-infecting Begomovirus in East Africa. Arch Virol 162(11):3439–3445 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Delphinium spp. (Larkspur) Family: Ranunculaceae

Aconitum latent virus Taxonomic position Genus: Carlavirus

Ornamental

(AcLV)

Family: Betaflexiviridae

AcLV infection in plants of Delphinium spp. was reported from Japan (Fuji et al. 2002). The virusinfected larkspur plants exhibit faint mosaic on leaves, with occasional spotting on flowers. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible through vegetative propagation. For more details of AcLV, refer to Aconitum spp.

D

856

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Delphinium spp. (Larkspur)

(ArMV)

Family: Secoviridae

ArMV infection in plants of Delphinium elatum was reported from the UK (Ahmed and Bailiss 1975). The virus was spreading through infected cuttings, but was not sap-transmissible to Delphinium. The virus is transmitted by the nematode vectors (Xiphinema spp.) in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Broad bean wilt virus Taxonomic position Genus: Fabavirus

(BBWV)

Family: Secoviridae

BBWV (but without specifying either of the currently recognized species BBWV-1 or BBWV-2) infection in plants of Delphinium elatum was reported from the UK (Ahmed and Bailiss 1975). The virus-infected larkspur plants exhibit symptoms of stunting and leaf deformation. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of BBWV, refer to Vicia faba.

Cherry leaf roll virus Taxonomic position Genus: Nepovirus

(CLRV)

Family: Secoviridae

CLRV infection in plants of Delphinium elatum was reported from the UK (Ahmed and Bailiss 1975). The virus-infected larkspur plants did not show any symptoms. The virus is transmitted by the nematode vectors, Xiphinema coxi, X. diversicaudatum, and X. vuittenezi, in a non-persistent manner. The virus is also transmissible by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants. For more details of CLRV, refer to Prunus avium.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Delphinium spp. was reported from the UK, Lithuania, Israel, California, and elsewhere in the USA (Severin and Dickson 1942; Ahmed and Bailiss 1975; Gera 1994; Chen et al. 1995; Samuitiene and Navalinskiene 2008). The virus-infected larkspur plants show mosaic pattern consisting of yellow spots of various shapes including ringspots (and initially described as Delphinium ringspot virus). The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Delphinium spp. (Larkspur)

857

Delphinium vein-clearing virus

(DeVCV)

Taxonomic position DeVCV is a tentative member of the genus Potyvirus and family Potivyridae Geographical distribution DeVCV infection in plants of wild larkspur (Delphinium glaucum) was reported from Alaska, USA (Robertson and Brown 2009). Symptoms and host(s) The virus-infected wild larkspur plants show obvious vein-clearing and chlorotic mosaic and in subsequent season were obviously stunted compared to plants without symptoms. Transmission No mechanical transmission was observed; as a potyvirus, aphid transmission is expected. Virion properties and genome The virions are flexuous filaments. The genome is a single molecule of positive-sense single-stranded RNA. A partial genome sequence of 1695 nt is available (FJ349327) (Robertson and Brown 2009; Wylie et al. 2017).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Tospovirus

(INSV)

Family: Peribunyaviridae

INSV infection in plants of Delphinium spp. was reported from Italy (Dellavalle et al. 1999). The virusinfected larkspur plants exhibit symptoms of mottling/mosaic, foliar distortion and chlorosis, and ringspots. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Potato virus X

(PVX)

Taxonomic position Genus: Potexvirus

Family: Alphaflexiviridae

PVX infection in plants of Delphinium elatum was reported from the UK (Ahmed and Bailiss 1975). The virus is transmissible by mechanical sap-inoculation, and also by contact between plants. For more details of PVX, refer to Solanum tuberosum.

Raspberry ringspot virus Taxonomic position Genus: Nepovirus

(RpRSV)

Family: Secoviridae

D

858

Delphinium spp. (Larkspur)

RpRSV infection in plants of Delphinium elatum was reported from the UK (Ahmed and Bailiss 1975). The virus is transmitted by the nematode vectors (Longidorus spp.) in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of RpRSV, refer to Rubus spp.

Strawberry latent ringspot virus Taxonomic position Genus: Unassigned

(SLRSV)

Family: Secoviridae

SLRSV infection in plants of Delphinium elatum was reported from the UK (Ahmed and Bailiss 1975). The virus is transmitted by the nematode vectors (Xiphinema spp.) and is also transmissible by mechanical sap-inoculation. The virus is transmissible by grafting. For more details of SLRSV, refer to Fragaria spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Tospovirus

(TSWV)

Family: Peribunyaviridae

TSWV infection in plants of Delphinium spp. was reported from California, USA, and the UK (Gardner et al. 1935; Smith 1937; Severin 1942; Hausbeck et al. 1992). The virus-infected larkspur plants exhibit symptoms of black rings or numerous double concentric rings, or patches of dead tissue may appear on the older leaves. The younger leaves may be malformed with edges yellow, necrotic, and inwardly curled. Necrotic patches may develop on the stems and older leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Ahmed AH, Bailiss KW (1975) Virus infection of delphinium in Britain. J Hortic Sci 50:47–54 Chen YK, Chen CC, Ko WF, Chen MJ (1995) Isolation and identification of cucumber mosaic virus from larkspur. Plant Protect Bull (Taipei) 37(3):319–330 Dellavalle G, Ciuffo M, Roggero P, Lisa V, Minuto A, Minuto G, Rapetti S (1999) Tospoviruses in Delphinium sp., Gazania, Marguerite, Celery, Tragopogon porrifolius and Solanum rantonnetti in Liguria (Northern Italy). Inf Fitopatol 49:63–64 Fuji S, Yamamoto H, Inoue M, Yamashita K, Fukui Y, Furuya H, Naito H (2002) Complete nucleotide sequence of the genomic RNA of Aconitum latent virus (genus Carlavirus) isolated from Delphinium sp. Arch Virol 147:865–870 Gardner MW, Tompkins CM, Whipple OC (1935) Spotted wilt of truck crops and ornamental plants. Phytopathology 25:17 Gera A (1994) The natural occurrence of Cucumber mosaic virus in ornamentals in Israel. Acta Hortic 377:99–106 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among green house ornamentals in Pennsylvania. Plant Dis 76:795–800 Robertson NL, Brown KL (2009) Identification and molecular characterization of a Potyvirus isolated from native larkspur (Delphinium glaucum) in Alaska. Plant Dis 93:428 Samuitiene M, Navalinskiene M (2008) Occurrence of Cucumber mosaic cucumovirus on ornamental plants in Lithuania. Zemdirbyste-Agriculture 95:135–143 Severin H (1942) The susceptibility of perennial delphiniums to six viruses. Hilgardia 14(10):549–570

Dendrobium spp.

859

Severin H, Dickson R (1942) Perennial-delphinium ringspot. Hilgardia 14(8):465–490 Smith KM (1937) A text book of plant virus diseases. Churchill, London Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354

Dendrobium spp.

D

Family: Orchidaceae

Ornamental

Bean common mosaic virus

(BCMV)

Synonyms Dendrobium mosaic virus (DeMV) Taxonomic position Genus: Potyvirus

Family: Potyviridae

DeMV was first reported from plants of Dendrobium spp. in Japan (Inouye 1973), and DeMV was characterized from D. superbum in Hawaii, USA (Hu et al. 1995). The virus-infected Dendrobium plants exhibit symptoms of chlorosis, mosaic and distortion of leaves, and color breaking and distortion of flowers (Hu et al. 1995). The virus is naturally transmitted by aphids (Myzus persicae) in a nonpersistent manner, and is also transmissible mechanically during orchid propagation. For more details of BCMV, refer to Phaseolus vulgaris.

Cucumber mosaic virus

(CMV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV infection in plants of Dendrobium spp. was reported from Japan, Israel, and Brazil (Inouye 1969; Lawson and Hsu 1995; Alexandre et al. 2005). The virus-infected dendrobium plants show mosaic, chlorosis, and leaf distortion symptoms. The virus is transmitted by a number of aphid species in a nonpersistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Cymbidium mosaic virus Taxonomic position Genus: Potexvirus

(CymMV)

Family: Alphaflexiviridae

CymMV infection reported in plants of Dendrobium orchids worldwide including Puerto Rico, Ecuador, Florida, Hawaii, Bangkok, and Thailand (Anamthawat and Vajrabhaya 1979; Hu et al.

860

Dendrobium spp.

1993; Elliott et al. 1996; Khentry et al. 2006a, b; Vaughan et al. 2008). The virus-infected dendrobium plants exhibit symptoms of chlorotic flecks and lesions in leaf tissues and necrosis in flowers. Although readily mechanically transmissible, no vector is known, and if any seed-transmission occurs, it must be at low efficiency (Yuen et al. 1979; Porter et al. 1996). For more details of CymMV, refer to Cymbidium spp.

Dendrobium leaf streak virus

(DLSV)

Synonyms Dendrobium rhabdovirus Taxonomic position DSLV is a tentative member of the family Rhabdoviridae Geographical distribution DLSV infection was first reported in plants of Dendrobium spp. in Germany (Petzold 1971; Duvel and Peters 1971). The virus spreads in the USA (Ali et al. 1974) and other locations. Symptoms and host(s) Typically, only foliar symptoms are associated with infection, including ringspots, except that white streaks were observed in flowers of Dendrobium phalaenopsis (Ali et al. 1974). Transmission An aerial vector is suspected, but not identified. Virion properties and genome Particles are reported to be c.180–320  85 nm and bacilliform or bullet-shaped (Ali et al. 1974). No sequence is currently available.

Dendrobium vein necrosis virus

(DVNV)

Taxonomic position DVNV is a tentative member of the genus Closterovirus and family Closteroviridae Geographical distribution DVNV was first reported in plants of Dendrobium phalaenopsis from Germany by Lesemann (1977). The virus spreads in the USA. Symptoms and host(s) The virus-infected dendrobium plants exhibit symptoms of vein necrosis in leaves, petals, and sepals. Transmission The virus is transmitted by aphid vectors (Zettler et al. 1990).

Dendrobium spp.

861

Virion properties and genome The virions are flexuous filaments with a clear modal length of 1865 nm and 10–12 nm wide. No sequence is currently available.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Tospovirus

(INSV)

D Family: Peribunyaviridae

INSV infection in plants of Dendrobium orchids was reported from China (Zhang et al. 2010). The virus-infected dendrobium plants exhibit symptoms of large chlorotic/necrotic ringspot symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Odontoglossum ringspot virus Taxonomic position Genus: Tobamovirus

(ORSV)

Family: Virgaviridae

ORSV infection in plants of Dendrobium spp. was reported from Hawaii, Thailand, and India (Hu et al. 1993; Khentry et al. 2006a, b; Sherpa et al. 2006). The virus-infected dendrobium plants exhibit symptoms of chlorotic and necrotic spots, streaks, and rings. The rings may be distinguished as ringspots with a distinct pinpoint spot surrounded by a circle of chlorotic tissue. The flowers from the infected plants are variegated. The normal pigment of the sepals and petals is replaced with irregular splotches of pigment which is more intense in color than that of normal flower. Individual flowers on the same plant may show different degrees of color breaking from one flowering to the next. The virus is transmitted by means not involving a vector. The virus is transmissible mechanically by sap and also by contact between plants. A contaminated knife used in propagation or cutting flowers will transmit viruses to healthy plants. For more details of ORSV, refer to Odontoglossum grande.

Rhopalanthe virus Y Taxonomic position Genus: Potyvirus

(RhVY)

Family: Potyviridae

Geographical distribution RhVY infection in plants of Dendrobium philippinense (Rhopalanthe) was reported from Australia (Gibbs et al. 2000). Transmission The virus is transmitted by aphid vectors in a non-persistent manner, and the virus is also mechanically sap-transmissible.

862

Dendrobium spp.

Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long and 11–13 nm wide. The genome is a single molecule of linear positive-sense single-stranded RNA. A partial genome sequence of 1622 nt is available (AF185956) (Revers and Garcia 2015; Wylie et al. 2017).

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV infection in plants of Dendrobium anosmum was reported from Hawaii (Watanabe et al. 2016). The virus-infected plants exhibit deformed flowers displaying color break and mild chlorotic streaking of the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of WMV, refer to Citrullus lanatus.

References Alexandre MAV, Rivas EB, Tozetto ARP, Duarte LML (2005) An annotated list on the natural occurrence of viruses in ornamental plants in Brazil. Instituto Biologico, Sao Paulo, 54 p Ali S, Lawson RH, Ishii M (1974) A bacilliform virus in white-streaked Dendrobium phalaenopsis flowers. Am Orchid Soc Bull 43:529–533 Anamthawat K, Vajrabhaya T (1979) Survey of the occurrence of cymbidium mosaic virus (CymMV) in Dendrobium orchid in Bangkok. Aust Orchid Rev 44(2):79–81 Duvel D, Peters KR (1971) Virusahnliche partikel in Dendrobium antennatum Ldl. Gartenwelt 71:52–54 Elliott MS, Zettler FW, Zimmerman MT, Barnett OW Jr, Le Grande MD (1996) Problems with interpretation of serological assays in a virus survey of orchid species from Puerto Rico, Ecuador, and Florida. Plant Dis 80:1160–1164 Gibbs A, Mackenzie A, Blanchfield A, Cross P, Wilson C, Kitajima E, Nightingale M, Clements M (2000) Viruses of orchids in Australia: their identification, biology and control. Aust Orchid Rev 65:10–21 Hu JS, Ferreira S, Wang M, Xu MQ (1993) Detection of cymbidium mosaic virus, odontoglossum ringspot virus, tomato spotted wilt virus, and potyviruses infecting orchids in Hawaii. Plant Dis 77:464–468 Hu JS, Ferreria S, Wang M, Borth WB, Mink G, Jordan R (1995) Purification, host range, serology, and partial sequencing of Dendrobium mosaic potyvirus, a new member of the bean common mosaic virus subgroup. Phytopathology 85:542–546 Inouye N (1969) Cucumber mosaic virus isolated from Dendrobium. Agric Res (Kurashiki) 53:49–60 Inouye N (1973) A new virus isolated from Dendrobium. Ann Phytopathol Soc Jpn 39:367–368 Khentry Y, Paradornuwat A, Tantiwiwat S, Phansiri S, Thaveechai N (2006a) Incidence of Cymbidium mosaic virus and Odontoglossum ringspot virus in Dendrobium spp. in Thailand. Crop Prot 25:926–932 Khentry Y, Paradornuwat A, Tantiwiwat S, Phansiri S, Thaveechai N (2006b) Incidence of Cymbidium mosaic virus and Odontoglossum ringspot virus on in vitro Thai native orchid seedlings and cultivated orchid mericlones. Kasetsart J (Nat Sci) 40:49–57 Lawson RH, Hsu HT (1995) Orchid. Chapter 36. In: Loebenstein G, Lawson RH, Brunt AA (eds) Virus and virus-like diseases of bulb and flower crops. Wiley, Chichester, pp 409–420 Lesemann DE (1977) Long filamentous virus like particles associated with vein necrosis of Dendrobium phalaenopsis. Phytopathol Z 89:330–339 Petzold H (1971) Der electroenmikroskopische nachweis eines bacilliforme virus on blattfleckernkranken Dendrobien. J Phytopathol 70:43–52 Porter KG, Kuehnle AR, Hu JS (1996) Lack of seed transmission of cymbidium mosaic virus in dendrobium. Lindleyana 11(4):211–213 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Sherpa AR, Bag TK, Hallan V, Zaidi AA (2006) Detection of Odontoglossum ringspot virus in orchids from Sikkim, India. Aust Plant Pathol 35:69–71 Vaughan SP, Grisoni M, Kumagai MH, Kuehnle AR (2008) Characterization of Hawaiian isolates of Cymbidium mosaic virus (CymMV) co-infecting Dendrobium orchid. Arch Virol 153:1185–1189

Deschampsia cespitosa (Tufted hair grass)

863

Watanabe S, Ruschel R, Marrero G, Sether D, Borth W, Hu J, Melzer M (2016) A distinct lineage of Watermelon mosaic virus naturally infects honohono orchid (Dendrobium anosmum) and passionfruit (Passiflora edulis) in Hawaii. New Dis Rep 34:13 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354 Yuen CKKH, Kamemoto H, Ishii M (1979) Transmission of Cymbidium mosaic virus through seed propagation in dendrobium. Am Orchid Soc Bull 48:1245–1247 Zettler FH, Ko NJ, Wisler GC, Eliote MS, Wong SM (1990) Viruses of orchids and their control. Plant Dis 74:621–626 Zhang Q, Ding Y-M, Li M (2010) First report of Impatiens necrotic spot virus infecting Phalaenopsis and Dendrobium orchids in Yunnan Province. China Plant Dis 94:915

Deschampsia antarctica (Antarctic hair grass) Family: Poaceae

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

CGMMV infection in plants of Deschampsia antarctica was reported from Argentina Islands, Antarctica (Polischuk et al. 2007). There is no known vector for this virus. The virus is transmissible by mechanical sap-inoculation, and also by contact between plants. For more details of CGMMV, refer to Cucumis sativus.

References Polischuk V, Budzanivska I, Shevchenko T, Oliynik S (2007) Evidence for plant viruses in the region of Argentina Islands, Antarctica. FEMS Microbiol Ecol 59:409–417

Deschampsia cespitosa (Tufted hair grass) Family: Poaceae

Ryegrass mosaic virus Taxonomic position Genus: Rymovirus

Weed host

(RGMV)

Family: Potyviridae

RGMV infection in plants of Deschampsia cespitosa was reported from Lithuania (Urbanavieiene 2004). The virus-infected tufted hair grass plants exhibit symptoms of chlorotic flecking, mosaic, and necrotic streaks. The virus is transmitted by the mite vector Abacarus hystrix in a semi-persistent

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Desmodium spp.

manner, and is also transmissible through mechanical sap-inoculation. For more details of RGMV, refer to Lolium spp.

References Urbanavieiene L (2004) Viral diseases of poaceae family plants. Pemes Ukio Mokslai 1:19–23

Desmodium spp. Family: Fabaceae

Ornamental

Cowpea chlorotic mottle virus Taxonomic position Genus: Bromovirus

(CCMV)

Family: Bromoviridae

CCMV infection in plants of Desmodium heterocarpon was reported from Nigeria (Walters and Dodd 1969; Thottappilly et al. 1993). The virus-infected desmodium plants exhibit bright yellow mosaic symptoms, which varied from small circular spots or stipples to large patches, occasionally covering the entire leaflet. The virus is transmitted by beetle vectors and is also transmissible by mechanical sapinoculation. For more details of CCMV, refer to Vigna unguiculata.

Cowpea severe mosaic virus Taxonomic position Genus: Comovirus

(CPSMV)

Family: Secoviridae

CPSMV infection in plants of Desmodium canescens was reported from Illinois (McLaughlin et al. 1978). The virus-infected desmodium plants induce systemic mottle or mosaic, frequently with severe blistering and distortion of leaflets. The virus is transmitted by the beetle vector, Cerotoma trifurcata, and is also transmissible by mechanical sap-inoculation. For more details of CPSMV, refer to Vigna unguiculata.

Desmodium leaf distortion virus Taxonomic position Genus: Begomovirus

(DesLDV)

Family: Geminiviridae

Geographical distribution DeLDV infection in plants of Desmodium glabrum was reported from Mexico (Hernandez-Zepeda et al. 2007, 2009; Fiallo-Olive et al. 2013).

Desmodium spp.

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Symptoms and host(s) The virus-infected desmodium plants show leaf distortion symptoms. Transmission The transmission of the DesLDV has not been investigated. It is likely that, in common with other begomoviruses, DesLDV is transmitted by the whitefly vector Bemisia tabaci in a circulative, nonpropagative manner.

D Virion properties and genome The virion structure of DesLDV has not been investigated. In common with all geminiviruses, the virions of DesLDV are likely geminate (twinned quasi-icosahedra). DesLDV is typical of the majority of begomoviruses native to the New World with a bipartite genome consisting of two circular, single-stranded DNA components. DNA-A contains 2569 nt (DQ875870; NC_008494) and DNA-B of 2514 nt (DQ875871 = NC_008495) (Briddon 2001; Hernandez-Zepeda et al. 2007, 2009; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the New World, the DNA-A component of DesLDVencodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for DesLDV.

Desmodium mottle virus Taxonomic position Genus: Begomovirus

(DesMoV)

Family: Geminiviridae

Geographical distribution DesMoV infection in plants of Desmodium spp. was reported from Uganda (Mollel et al. 2017). Symptoms and host(s) The virus-infected desmodium plants exhibit mottling symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is bipartite consisting of two circular, single stranded DNA components. DNA-A contains 2767 nt (KY294725 = NC_039001, KY294724) and DNA-B of 2713 nt (KY294727, KY924726) (Briddon 2001; Brown et al. 2015; Mollel et al. 2017; Zerbini et al. 2017).

Desmodium yellow mottle virus Taxonomic position Genus: Tymovirus

(DYMoV)

Family: Tymoviridae

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Desmodium spp.

Geographical distribution DYMoV infection in plants of Desmodium spp. was reported from Arkansas in the USA (Walters and Scott 1972; Larson et al. 2000). Symptoms and host(s) The virus-infected desmodium plants exhibit yellow mottle and leaf malformation symptoms. Transmission The virus is transmitted by beetle vectors in a semi-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, isometric capsids with T = 3 icosahedral symmetry, about 30 nm in diameter. The genome consists of a single molecule of positive-sense, ssRNA. The 30 -terminus has a tRNA-like structure. A partial genome sequence of 688 nt is available (AF035201) (Larson et al. 2000; Martelli et al. 2002).

Macroptilium yellow spot virus Taxonomic position Genus: Begomovirus

(MacYSV)

Family: Geminiviridae

MacYSV infection in plants of Desmodium glabrum was reported from Brazil (Fontenele et al. 2016). The virus-infected desmodium plants exhibit yellow mosaic, leaf curling, rugosity, and leaf deformation symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative nonpropagative manner. For more details of MacYSV, refer to Macroptilium spp.

Peanut mottle virus Taxonomic position Genus: Potyvirus

(PeMoV)

Family: Potyviridae

PeMoV infection in plants of Desmodium canum was reported from the USA (Demski et al. 1981). The virus-infected desmodium plants exhibit bright chlorotic mottle symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of PeMoV, refer to Arachis hypogaea.

Peanut stunt virus Taxonomic position Genus: Cucumovirus

(PSV)

Family: Bromoviridae

PSV infection in plants of Desmodium spp. was reported from Brazil (Gillaspie and Ghabrial 1998). The virus-infected desmodium plants exhibit mild mosaic symptoms. The virus is transmitted by aphid

Desmodium spp.

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vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of PSV, refer to Arachis hypogaea.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Tospovirus

(TSWV)

Family: Peribunyaviridae

TSWV infection in plants of Desmodium spp. was reported from Hawaii and Georgia (Cho et al. 1987; Mullis et al. 2009). The virus-infected desmodium plants exhibit vein yellowing and streaking symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Cho JJ, Mau RFL, Mitchell WC, Gonsalves D, Yudin LS (1987) Host list of plants susceptible to tomato spotted wilt virus (TSWV), Research Extension Series; RES-078. University of Hawaii, Honolulu, 10 p Demski J, Khan MA, Wells HD, Miller JD (1981) Peanut mottle virus in forage legumes. Plant Dis 65:359–362 Fiallo-Olive E, Chirinos DT, Geraud-Pouey F, Moriones E, Navas-Castillo J (2013) Complete genome sequences of two begomoviruses infecting weeds in Venezuela. Arch Virol 158(1):277–280 Fontenele MRS, Poppiel R, Lessa Matos VOR, Costa AF, Faria JC, Ribeiro SG (2016) First report of Macroptillium yellow spot virus in Desmodium glabrum in Brazil. Plant Dis 100:657 Gillaspie AG Jr, Ghabrial SA (1998) First report of Peanut stunt cucumovirus naturally infecting Desmodium sp. Plant Dis 82:1402 Hernandez-Zepeda C, Idris AM, Carnevali G, Brown JK, Moreno-Valenzuela OA (2007) Molecular characterization and phylogenetic relationships of two new bipartite begomovirus infecting malvaceous plants in Yucatan, Mexico. Virus Genes 35(2):369–377 Hernandez-Zepeda C, Argüello-Astorga G, Idris AM, Carnevali G, Brown JK, Moreno-Valenzuela OA (2009) Molecular characterization and phylogenetic relationships of Desmodium leaf distortion virus (DeLDV): a new begomovirus infecting Desmodium glabrum in Yucatan, Mexico. Virus Genes 39:371–374 Larson SB, Day J, Canady MA, Greenwood A, McPherson A (2000) Refined structure of desmodium yellow mottle tymovirus at 2.7 A resolution. J Mol Biol 301(3):625–642 Martelli GP, Sabanadzovic S, Abou-Ghanem Sabanadzovic N, Edwards MC, Dreher T (2002) The family Tymoviridae. Arch Virol 147(9):1837–1846 McLaughlin MR, Thongmeearkom P, Goodman RM, Milbrath GM, Ries SM, Royse DJ (1978) Isolation and beetle transmission of cowpea mosaic virus (severe subgroup) from Desmodium canescens and soybeans in Illinois. Plant Dis Reptr 62:1069–1073 Mollel HG, Sseruwagi P, Ndunguru J, Alicai T, Colvin J, Navas-Castillo J, Fiallo-Olive E (2017) Desmodium mottle virus, the first legumovirus (genus Begomovirus) from East Africa. Arch Virol 162:1799–1803 Mullis S, Bertrand PF, Brown SL, Csinos A, Diaz-Perez JC, Gitaitis R, Hickman LL, Johnson A, LaHue SS, Martinez N, McPherson RM, Nischwitz C, Reay-Jones F, Riley D, Sherwood J, Stevenson K, Wells L (2009) Tospoviruses in Solanaceae and other crops in the coastal plain of Georgia. University of Georgia, College of Agricultural and Environmental Sciences, Bulletin 1354, pp 51 Thottappilly G, Sehgal OP, Rossel HW (1993) Characteristics of a cowpea chlorotic mottle virus isolate from Nigeria. Plant Dis 77:60–63

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Dianthus amurensis (Amur pink)

Walters HJ, Dodd NL (1969) Identification and beetle transmission of an isolate of Cowpea chlorotic mottle virus from Desmodium. Phytopathology 59:1055 Walters HJ, Scott HA (1972) Host range and some properties of Desmodium yellow mottle virus. Phytopathology 62:125–128 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

Dianthus amurensis (Amur pink) Family: Caryophyllaceae

Medicinal

Carnation vein mottle virus Taxonomic position Genus: Potyvirus

(CVMoV)

Family: Potyviridae

CVMoV infection in plants of Dianthus amurensis was reported from China (Li et al. 2014). The virusinfected Amur pink plants exhibit mosaic, leaf cupping, leaf distortion, reduction in leaf size, and flower-breaking symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible through mechanical sap-inoculation. For more details of CVMoV, refer to Dianthus caryophyllus.

References Li MR, Zhu J, Gao J, Li RH, Li F (2014) First report of carnation vein mottle virus infecting Dianthus amurensis in China. Plant Dis 98:1747

Dianthus barbatus (Sweet William) Family: Caryophyllaceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

Family: Secoviridae

ArMV infection in plants of Dianthus barbatus was reported from Lithuania (Samuitiene et al. 2008). The virus-infected sweet William plants exhibit symptoms of necrosis, stunting, and death of the plants. The virus is transmitted by the nematode vectors (Xiphinema spp.) in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Dianthus barbatus (Sweet William)

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Carnation etched ring virus Taxonomic position Genus: Caulimovirus

(CERV)

Family: Caulimoviridae

CERV infection in plants of Dianthus barbatus was reported from the UK and Iran (Hollings and Stone 1961; Rubio-Huertos et al. 1972; Ashnayi et al. 2012). The virus is transmitted by the aphid vector, Myzus persicae, in a semi-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CERV, refer to Dianthus caryophyllus.

Carnation latent virus Taxonomic position Genus: Carlavirus

(CLV)

Family: Betaflexiviridae

CLV infection in plants of Dianthus barbatus was reported from the UK (Kassanis 1954). The virusinfected sweet William plants are typically symptomless. The virus is transmitted by the aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of CLV, refer to Dianthus caryophyllus.

Carnation mottle virus

(CarMV)

Taxonomic position Genus: Alphacarmovirus

Family: Tombusviridae

CarMV was first reported in the UK on Dianthus spp. by Kassanis (1955). The virus infection in Dianthus barbatus is generally symptomless. Hollings and Stone (1964) have reported flower break symptoms consisting of narrow pink lines on red-flowered clones. The virus is transmitted by means not involving a vector. The virus is transmissible by mechanical sap-inoculation, by grafting, and also by contact between plants. For more details of CarMV, refer to Dianthus caryophyllus.

Carnation vein mottle virus Taxonomic position Genus: Potyvirus

(CVMoV)

Family: Potyviridae

CVMoV infection in plants of Dianthus barbatus was reported from Britain and Czechoslovakia (Mokra and Gotzova 1994). The virus-infected sweet William plants express symptoms of veinclearing followed by conspicuous light and green leaf mottle, flecks, rings, and spots on the lateral veins. Dwarfing of the infected plants and flower break in red- or pink-flowered varieties were also recorded. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible through mechanical sap-inoculation. For more details of CVMoV, refer to Dianthus caryophyllus.

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Dianthus barbatus (Sweet William)

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV was reported in India on Dianthus barbatus plants by Raj et al. (1993). The Indian strain of CMVCR is placed in CMV subgroup of 1 on the basis of sequence homology (Raj et al. 1995). The virusinfected sweet William plants show stunting and leaf crinkling symptoms. A number of aphid vectors transmit this virus in a non-persistent manner. The virus is also mechanically sap-transmissible to healthy D. barbatus and Vaccaria pyramidata (Caryophyllaceae) and to other herbaceous test plants. For more details of CMV, refer to Cucumis sativus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Tospovirus

(INSV)

Family: Peribunyaviridae

INSV infection in plants of Dianthus spp. was reported from Italy and Iran (Roggero et al. 1999; Ghotbi et al. 2005; Ghotbi 2013). The virus-infected sweet William plants exhibit symptoms of necrotic leaf spot, chlorosis, and stunting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Moroccan pepper virus Taxonomic position Genus: Tombusvirus

(MPV)

Family: Tombusviridae

MPV infection in plants of Dianthus barbatus was reported from Southern Iran (Alemzadeh and Ghorbani 2016). The virus-infected sweet William plants exhibit mosaic and ringspot symptoms. The virus is transmissible by mechanical inoculation to three to nine families and by grafting. No vector is known for this virus. For more details of MPV, refer to Capsicum annuum.

Pelargonium leaf curl virus Taxonomic position Genus: Tombusvirus

(PLCV)

Family: Tombusviridae

PLCV infection in plants of Dianthus barbatus was reported from Southern Iran (Alemzadeh and Ghorbani 2016). The virus-infected sweet William plants exhibit leaf curling symptoms. No vector is reported for this virus. The virus is mechanically sap-transmissible. For more details of PLCV, refer to Pelargonium spp.

Dianthus caryophyllus (Carnation)

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References Alemzadeh E, Ghorbani A (2016) Occurrence of Pelargonium leaf curl virus and Moroccan pepper virus on natural hosts. Aust Plant Dis Notes 11:8 Ashnayi M, Jafarpour B, Malekzadeh Shafarudi S, Mirshamsi Kakhki A, Heydariniya Z (2012) Detection of carnation etched ring virus in greenhouses of Khorasan Razavi and northern Khorasan provinces by ELISA and PCR. J Plant Prot (Ag Sci Technol) 2:171–177 Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iran J Plant Pathol 49:41 Ghotbi T, Sharaeen N, Winter S (2005) Occurrence of tospoviruses in ornamental and weed species in Markazi and Tehran provinces in Iran. Plant Dis 89:425–429 Hollings M, Stone OM (1961) Carnation etched ring: a preliminary report on an undescribed disease. Rep Glasshouse Crop Res Inst for 1960:94–95 Hollings M, Stone OM (1964) Investigation on carnation viruses. I. Carnation mottle. Ann Appl Biol 53:103–118 Kassanis B (1954) A virus latent in carnation and potato plants. Nature 173:1097–1098 Kassanis V (1955) Some properties of four viruses isolated from carnation plants. Ann Appl Biol 43:103–113 Mokra V, Gotzova B (1994) Detection of carnation vein mottle virus in Dianthus barbatus. Acta Hortic 377:363–365 Raj SK, Aminuddin, Srivastava KM, Singh BP (1993) Natural infection of Cucumber mosaic virus on Dianthus barbatus in India. Plant Pathol 42:811–813 Raj SK, Haq QMR, Srivastava KM, Singh BP (1995) Sequence homology of two Indian isolates of CMV at N-terminal amino acid sequences of the CP gene with CMV strains. J Plant Biochem Biotechnol 4:77–80 Roggero P, Ciuffo M, Dellavalle G, Gotta P, Gallo S, Peters D (1999) Additional ornamental species as hosts of impatiens necrotic spot virus in Italy. Plant Dis 83:967 Rubio-Huertos M, Castro S, Fujisawa I, Matsui C (1972) Electron microscopy of the formation of Carnation etched ring virus intracellular inclusion bodies. J Gen Virol 15:257–260 Samuitiene M, Navalinskiene M, Jackeviciene E (2008) Arabis mosaic virus on ornamental plants. Biologia 54:264–268

Dianthus caryophyllus (Carnation) Family: Caryophyllaceae

Ornamental

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

ArMV infection in plants of Dianthus caryophyllus has been reported in the Netherlands, and New Zealand (Bennet and Milne 1976; Loebenstein et al. 1995). The virus-infected carnation plants show shortened internodes with prolific lateral branching, giving the plants a bushy appearance. The flowering was extremely retarded or absent. The virus is transmitted by the nematode vector Xiphinema diversicaudatum in a non-persistent manner, and is also transmissible by mechanical sap-inoculation (Hakkaart et al. 1972). For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Carnation bacilliform virus

(CBV)

Taxonomic position CBV is a tentative member of the family Rhabdoviridae.

D

872

Dianthus caryophyllus (Carnation)

Geographical distribution CBV infection in plants of Dianthus caryophyllus was reported from New Zealand (Bennet and Milne 1976). Virion properties and genome The virions are bullet-shaped and measure 55 nm in diameter and 260 nm in length. The genome is not segmented and contains a single molecule of linear single-stranded RNA.

Carnation cryptic virus 1 Taxonomic position Genus: Unassigned

(CCV1)

Family: Partitiviridae

Geographical distribution CCV1 is probably distributed worldwide wherever Dianthus caryophyllus plants are grown (Lisa et al. 1980, 1981). Symptoms and host(s) The virus-infected carnation plants do not exhibit any symptoms. The virus was found to persist in Dianthus caryophyllus, D. superbus, D. carthusianorum, and D. gratianopolitanus maintained in tissue culture for several years; in some plants one to three additional dsRNA species of 1100–3600 bp were also detected (Szego et al. 2006). Transmission The virus is transmitted by means not involving a vector. The virus is not transmissible by mechanical inoculation, not transmissible by grafting, not transmissible by contact between plants, and transmitted by seed (probably, and also in cuttings). Virion properties and genome The virions are isometric, non-enveloped, and 29 nm in diameter. The genome consists of two dsRNA segments (Vainio et al. 2018).

Carnation etched ring virus Taxonomic position Genus: Caulimovirus

(CERV)

Family: Caulimoviridae

Geographical distribution CERV infection in plants of Dianthus caryophyllus is probably distributed worldwide (Bennet and Milne 1976; McRitchie 1980; Rana 1984; Raikhy et al. 2003b; De la Torre-Almaraz et al. 2015). Symptoms and host(s) On many carnation varieties, the virus exhibits necrotic flecks, rings, and line patterns “etched” on leaves, while some other cultivars are symptomless.

Dianthus caryophyllus (Carnation)

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Transmission The virus is transmitted by the aphid vector, Myzus persicae, in a semi-persistent manner. The virus is lost by the vector when it moults and does not multiply in the vector, and is not transmitted congenitally to the progeny of the vector. The virus is transmissible by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants and not transmitted by seed. Virion properties and genome The virions are isometric, non-enveloped, and 45 nm in diameter with no obvious surface structure. The genome is a monopartite, circular double-stranded DNA of 7932 bp (NC_003498, X04658) (Fujisawa et al. 1972; Palkovics and Balazs 1996; Raikhy et al. 2006c; Hohn 2011).

Carnation Italian ringspot virus Taxonomic position Genus: Tombusvirus

(CIRV)

Family: Tombusviridae

Geographical distribution CIRV was first reported in plants of Dianthus caryophyllus in the UK in plants received from Italy and the USA (Hollings et al. 1970). The virus was also reported from Germany, Italy, Spain, and Korea (Buttner et al. 1987; Sanchez-Navarro et al. 1999). Symptoms and host(s) The virus-infected carnation plants exhibit transient chlorotic spots and rings in leaves and slight stunting. Transmission The virus was mechanically sap-transmissible to 62 of 104 plants tested. The use of virus-infected propagating material will spread the disease. The virus is transmissible by grafting. Virion properties and genome The virions are isometric and icosahedral (T = 3), non-enveloped, and about 30 nm in diameter. The genome is a monopartite, linear, positive-sense, single-stranded RNA of 4760 nt (NC_003500 = X85215) (Hollings et al. 1970; Rubino et al. 1995; White 2011).

Carnation latent virus Taxonomic position Genus: Carlavirus

(CLV)

Family: Betaflexiviridae

Geographical distribution CLV is probably distributed worldwide wherever Dianthus caryophyllus plants are grown (Bennet and Milne 1976; Sanchez-Navarro et al. 1999; Manisha et al. 2002). Symptoms and host(s) The virus-infected carnation plants do not show specific symptoms, but infected plants are less vigorous and produce fewer good flowers (Wetter 1971).

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874

Dianthus caryophyllus (Carnation)

Transmission The virus is transmitted by the aphid vector, Myzus persicae, in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation to a limited number of host plants belonging to the families Caryophyllaceae, Chenopodiaceae, and Solanaceae. The virus is not transmissible by contact between plants and not transmitted by seed. Virion properties and genome The virions are filaments, non-enveloped, and usually straight (or slightly curved) with a clear modal length of 650 nm and 12 nm wide (Brandes et al. 1959). The genome is single-stranded RNA of c.8.5–9 kb; partial genome sequences are available (X52627, AJ010697, X55897; FJ555525) (Meehan and Mills 1991). The genome comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Adams et al. 2004).

Carnation mottle virus Taxonomic position Genus: Alphacarmovirus

(CarMV)

Family: Tombusviridae

Geographical distribution CarMV is probably distributed worldwide wherever Dianthus caryophyllus plants are grown (Bennet and Milne 1976; Bremer 1978; Gallardo and Alvarez 1979; Bezic et al. 1983; Luisoni et al. 1984; Datta Gupta and Verma 1990; Sanchez-Navarro et al. 1999; Singh et al. 2005; Raikhy et al. 2006b; Safari et al. 2009; Cevik et al. 2010; De la Torre-Almaraz et al. 2015). Symptoms and host(s) The virus-infected carnation plants exhibit generally indistinct symptoms consisting of a mild mottling of the leaves, the symptoms are masked at the flowering stage. The virus infection reduce the number, size, and quality of flowers (Rana and Castellano 1984). Transmission The virus is transmitted by means not involving a vector. The virus is transmissible by mechanical sapinoculation, by grafting, and also by contact between plants (Chen et al. 2003). Virion properties and genome The virions are isometric 30 nm in diameter and have T = 3 icosahedral symmetry. The genome consists of a single molecule of linear positive-sense ssRNA of 4003 nt (AF192772 = NC_001265) (Guilley et al. 1985).

Carnation necrotic fleck virus

(CNFV)

Synonyms Carnation yellow fleck virus (CYFV) Taxonomic position Genus: Closterovirus

Family: Closteroviridae

Dianthus caryophyllus (Carnation)

875

Geographical distribution CNFV has probably been distributed worldwide wherever Dianthus caryophyllus plants are grown (Inouye and Mitsuhata 1973; Smookler and Loebenstein 1974; Bennet and Milne 1976; McRitchie and Wisler 1980; Batlle and Lavina 1994; Raikhy et al. 2003a; Karasev et al. 2008; Jordan et al. 2012). Symptoms and host(s) The virus-infected carnation plants exhibit whitish or necrotic streaks and reddish-purple flecks and spots on leaves and stems and complete necrosis on older leaves. Mayhew (1979) observed in California that infected carnation plants grown at low temperature had mild or no symptoms and disease severity increased with temperature (Smookler and Loebenstein 1974). Transmission The virus is transmitted by the aphid vector, Myzus persicae, in a semi-persistent manner. The virus is lost by the vector when it moults and does not multiply in the vector; nor is it transmitted congenitally to the progeny of the vector. The virus is transmissible by mechanical sap-inoculation (with difficulty) and not transmissible by contact between plants. Virion properties and genome The virions are filaments, non-enveloped, and usually flexuous with no clear modal length of 1400–1500 nm and 12 nm wide. The genome is a monopartite, linear, positive-sense, single-stranded RNA of 10,557 nt partial sequence and is available (GU234166) (Bar-Joseph and Smookler 1976; Agranovsky and Lesemann 2011). A complete genome of an isolate referred to as Carnation yellow fleck virus is 15,602 nt (GU234167 = NC_022978), but may represent a distinct virus (Karasev et al. unpublished - GU234167).

Carnation ringspot virus Taxonomic position Genus: Dianthovirus

(CRSV)

Family: Tombusviridae

Geographical distribution CRSV infection in plants of Dianthus caryophyllus was first reported from the UK by Kassanis (1955). The virus is probably distributed worldwide wherever carnations are grown (Bennet and Milne 1976; Magid and Robb 1990; Sanchez-Navarro et al. 1999; Raikhy et al. 2006a). Symptoms and host(s) The virus-infected carnation plants show rings, line patterns, and mottling on leaves, which also show lateral bending and buckling. The plants are stunted, flowers are smaller: bleaching, color breaking, and petal crinkle may also occur (Tremaine and Dodds 1985). Transmission Early reports indicated that CRSV is transmitted by the nematode vectors Longidorus elongatus, L. macrosoma, and Xiphinema diversicaudatum, but later reports suggest passive movement from roots to soil and uptake from contaminated soil via roots (Brown and Trudgill 1984). The virus is transmissible easily by mechanical inoculation to a number of fruit trees and weed hosts, transmissible by contact between plants, but is not transmitted by seed (Magid and Robb 1990).

D

876

Dianthus caryophyllus (Carnation)

Virion properties and genome The virions are isometric, non-enveloped, and 31–34 nm in diameter and have T = 3 icosahedral symmetry. The genome is a bipartite, linear, positive-sense, single-stranded RNA. RNA1 consists of 3840 nt (L18870 = NC_003530) and RNA2 of 1403 nt (M88589 = NC_003531) (Dodds et al. 1977; Hiruki 1987; Kendall and Lommel 1992; Ryabov et al. 1994).

Carnation vein mottle virus Taxonomic position Genus: Potyvirus

(CVMoV)

Family: Potyviridae

Geographical distribution The CVMoV is distributed wherever Dianthus caryophyllus plants are grown (Bennet and Milne 1976; Bezic et al. 1983; Stefanac and Wrischer 1983; Raj et al. 1990; Sanchez-Navarro et al. 1999; Raikhy et al. 2006a; Bayat and Ghotbi 2010; Harju et al. 2011). Symptoms and host(s) The virus-infected carnation plants exhibit symptoms of chlorotic mottle or green vein mottle on leaves and flower breaking in susceptible cultivars resulting in decreased flower yield. Transmission The virus is transmitted by an aphid vector primarily Myzus persicae in a non-persistent manner. The virus is also transmissible by mechanical sap-inoculation but has a restricted host range. The virus is not transmissible by contact between plants and not transmitted by seed (Hollings et al. 1977). Virion properties and genome The virions are non-enveloped, filaments usually flexuous of 790 nm in length and 12 nm width. The genome consists of a single molecule of linear, positive-sense ssRNA (Raj et al. 1990). A partial genome sequence of 2326 nt (AB017630) and 2704 nt (LC381969) are available (Sasaya et al. 2000; Revers and Garcia 2015; Wylie et al. 2017).

Carnation yellow stripe virus

(CYSV)

Taxonomic position CYSV is a tentative member of the genus Necrovirus Geographical distribution CYSV was first reported in plants of Dianthus caryophyllus from Apulia, Southern Italy (Gallitelli et al. 1979). Symptoms and host(s) The virus-infected carnation plants exhibit bright yellow stripes on the leaves (Gallitelli et al. 1979). Transmission The virus is transmissible by mechanical sap-inoculation, and several plants from (three to nine) families are susceptible.

Dianthus caryophyllus (Carnation)

877

Virion properties and genome The virions are isometric and 30 nm in diameter. The genome consists of single-stranded RNA. No sequence is available.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

D Family: Bromoviridae

CMV infection in plants of Dianthus caryophyllus was reported from Yugoslavia and Mexico (Bezic et al. 1983; Stefanac and Wrischer 1983; De la Torre-Almaraz et al. 2016). The virus-infected carnation plants show green mosaic and severe leaf deformation symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Eggplant mottled crinkle virus

(EMCV)

Synonyms Lisianthus necrosis virus (LNV)

Taxonomic position Genus: Tombusvirus

Family: Tombusviridae

LNV was reported in plants of Dianthus caryophyllus for the first time from Taiwan (Chen and Hsu 2002; Chen et al. 2002). The virus-infected carnation plants initially show disease symptoms on the upper leaves as numerous yellow spots that enlarge and fuse into large chlorotic patches and expand to cover the entire leaves, which eventually become necrotic. No vector is reported for this virus. The virus is mechanically sap-transmissible. For more details of EMCV, refer to Solanum melongena.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Tospovirus

(INSV)

Family: Peribunyaviridae

INSV infection in plants of Dianthus caryophyllus was reported from Italy and Iran (Roggero et al. 1999; Shahraeen et al. 2002). The virus-infected carnation plants show stunting, mosaic, and leaf malformation, and some plants had symptomless infections. The virus is transmitted by a thrips vector, Frankliniella occidentalis, in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

878

Dianthus caryophyllus (Carnation)

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Agranovsky AA, Lesemann D-E (2011) Closterovirus. Closteroviridae. In: The Springer index of viruses. Springer, New York, pp 327–333. https://doi.org/10.1007/978-0-387-95919-1_50 Bar-Joseph M, Smookler M (1976) Purification, properties, and serology of carnation yellow fleck virus. Phytopathology 66:835–838 Batlle A, Lavina A (1994) First report of Carnation necrotic fleck virus in Spain. Plant Dis 78(11):1122 Bayat H, Ghotbi T (2010) First report of Carnation vein mottle virus (CVMV) infecting Carnation in Iran. Iran J Plant Pathol 46(1):29 Bennet PR, Milne KS (1976) Carnation viruses in New Zealand. Acta Hortic 59:61–62 Bezic N, Štefanac Z, Miličić D, Wrischer M (1983) Occurrence of carnation vein mottle and cucumber mosaic viruses on carnations in Yugoslavia. Acta Bot Croat 42:21–27 Brandes J, Wetter C, Bagnall RH, Larson RH (1959) Size and shape of the particles of Potato virus S, Potato virus M, and Carnation latent virus. Phytopathology 49:443–446 Bremer K (1978) The carnation mottle virus in Finnish carnations. Ann Agric Fenn 17(1):36–38 Brown DJF, Trudgill DL (1984) The spread of carnation ringspot virus in soil with or without nematodes. Nematologica 30:102–104 Buttner C, Jacobi V, Koenig R (1987) Isolation of Carnation Italian ring spot virus from a creek in a forested area South West of Bonn. J Phytopathol 118:131–134 Cevik B, Bakir T, Koca G (2010) First report of Carnation mottle virus in Turkey. Plant Pathol 59:394 Chen CC, Hsu HT (2002) Occurrence of a severe strain of Lisianthus necrosis virus in imported carnation seedlings in Taiwan. Plant Dis 86(4):444 Chen CC, Chen YK, Ko WF, Hsu HT (2002) Characterization of Lisianthus necrosis virus (genus Necrovirus) isolated from Dianthus caryophyllus. Plant Pathol Bull 11:137–146 Chen CC, Lin CY, Ko WF, Jan FJ (2003) Isolation and characterization of Carnation mottle virus from carnation. Plant Pathol Bull 12:199–208 Datta Gupta M, Verma VS (1990) In vitro production of virus free plants from carnation cultivar dwarf fragrance rose infected with Carnation mottel virus. Int Nat J Trop Plant Dis 8:51–59 De la Torre-Almaraz R, Pallás V, Sánchez-Navarro J (2015) First report of Carnation mottle virus (CarMV) and Carnation etched ring virus (CERV) in carnation from Mexico. Plant Dis 99:1191 De la Torre-Almaraz R, Pallás V, Sánchez-Navarro J (2016) First report ofCucumber mosaic virus (CMV) and CARNA-5 in carnation in Mexico. Plant Dis 100:1509 Dodds JA, Tremaine JH, Ronald WP (1977) Some properties of carnation ringspot virus single- and double-stranded ribonucleic acid. Virology 83:322–328 Fujisawa I, Rubio-Huertos M, Matsui C (1972) Deoxyribonuclease digestion of the nucleic acid from carnation etched ring virus. Phytopathology 62:810–811 Gallardo M, Alvarez A (1979) Identification of carnation mottle virus in Chile. Agric Técnica, Chile 39(3):103–107 Gallitelli D, Castellano MA, Di Franco A, Rana GL (1979) Properties of carnation yellow stripe virus, a member of the tobacco necrosis virus group. Phytopathol Mediterr 18:31–40 Guilley H, Carrington JC, Balazs E, Jonard G, Richards K, Morris TJ (1985) Nucleotide sequence and genome organization of carnation mottle virus RNA. Nucleic Acids Res 13(18):6663–6677 Hakkaart FA, Van Hoof HA, Maat DZ (1972) A case of Arabis mosaic virus in carnations. Neth J Plant Pathol 78:15–18 Harju VA, Skelton AL, Monger WA, Forde SMD, Daly M, Nixon T, Lawson R, Bennett S, Mumford RA (2011) A report of viruses, viroids and phytoplasmas found in ornamental plants from 1999–2007: findings from Central Science Laboratory, UK. Acta Hortic 901:223–229 Hiruki C (1987) The dianthoviruses: a distinct group of isometric plant viruses with bipartite genome. Adv Virus Res 33:257–300 Hohn T (2011) Caulimovirus. Caulimoviridae. In: The Springer index of viruses. Springer, New York, pp 271–277. https://doi.org/10.1007/978-0-387-95919-1_41 Hollings M, Stone OM, Bouttell GC (1970) Carnation Italian ringspot virus. Ann Appl Biol 65:299–309 Hollings M, Stone OM, Atkey PT, Barton RJ (1977) Investigations on Carnation viruses. IV Carnation vein mottle virus. Ann Appl Biol 85:59–70 Inouye T, Mitsuhata K (1973) Carnation necrotic fleck virus. Ber Ohara Inst, Okayama Univ 15:195–205 Jordan R, Guaragna MA, Dolja V, Karasev A (2012) Detection and molecular characterization of two distinct closteroviruses infecting carnation. The 13th international symposium on virus diseases of ornamental plants Norway, 24–29th June 2012. p 16

Dianthus caryophyllus (Carnation)

879

Karasev AV, Dolja VV, Guaragna MA, Jordan R (2008) Two viruses are associated with carnation necrotic fleck disease. Phytopathology 98:S78 Kassanis B (1955) Some properties of four viruses isolated from carnation plants. Ann Appl Biol 43:103–113 Kendall TL, Lommel SA (1992) Nucleotide sequence of carnation ringspot dianthovirus RNA-2. J Gen Virol 73(9):2479–2482 Lisa V, Luisoni E, Milne RG (1980) Carnation cryptic virus. Acta Hortic 110:175–176 Lisa V, Boccardo G, Milne R (1981) Double-stranded ribonucleic acid from carnation cryptic virus. Virology 115:410–413 Loebenstein G, Lawson RH, Brunt AA (1995) Virus and virus-like diseases of bulb and flowering crops. Wiley, Chichester, p 543 Luisoni E, Lisa V, Roggero P (1984) Carnation mottle virus in Italy: Rilevamento e prevenzione delle forme normale e attenuate. Atti Giorgio Fitopatol 3:427–436 Magid AMA, Robb SM (1990) Investigations on viruses of pinks (Dianthus sp.) in the United Kingdom. Arab J Plant Protect 8:121–126 Manisha M, Bhardwaj SV, Mangal AK, Kaur R (2002) Production of Carnation latent virus free stock of carnations. Phytomorphology 52:1–6 Mayhew DE (1979) Carnation necrotic fleck virus in California. Plant Dis Reptr 63:978–980 McRitchie JJ (1980) Carnation etched ring. Plant Pathology Circular No 211. Fla Dept Agric and Consumer Serv Division of Plant Industry. Gainesville, Florida, USA. McRitchie JJ, Wisler GC (1980) Carnation necrotic fleck. Plant Pathology Circular No 213. Fla Dept Agric and Consumer Serv Division of Plant Industry. Florida, USA. Meehan BM, Mills PR (1991) Nucleotide sequence of the 30 -terminal region of carnation latent virus. Intervirology 32(4):262–267 Palkovics L, Balazs E (1996) Simple and rapid detection of carnation etched ring virus by polymerase chain reaction. Acta Phytopathol Entomol Hung 31:161–168 Raikhy G, Hallan V, Kulshrestha S, Sharma ML, Ram R, Zaidi AA (2003a) First report of Carnation necrotic fleck virus (CNFV) infecting carnations in India. Plant Pathol 52:801 Raikhy G, Hallan V, Kulshrestha S, Lal Sharma M, Ram R, Zaidi AA (2003b) Molecular characterization of a Carnation etched ring virus isolate from India. Acta Virol 47:105–111 Raikhy G, Hallan V, Kulshrestha S, Ram R, Zaidi AA (2006a) Detection of Carnation ringspot and Carnation vein mottle virus (es) in carnation cultivars in India. Acta Hortic 722:247–258 Raikhy G, Hallan V, Kulshrestha S, Ram R, Zaidi AA (2006b) Multiplex PCR and genome analysis of Carnation mottle virus Indian isolate. Curr Sci 90:74–82 Raikhy G, Hallan V, Kulshrestha S, Ram R, Zaidi AA (2006c) Complete nucleotide sequence of an Indian isolate of Carnation etched ring virus and its homology with other Caulimoviruses. Curr Sci 90:176–187 Raj SK, Aslam M, Singh BP (1990) Purification and some properties of a virus causing chlorotic mottle disease in carnations (Dianthus caryophyllus L). Curr Sci 59:997–999 Rana GL (1984) Presence of Carnation etched ring virus in Italy. Informatore Fitopatologico 34:51–53 Rana GL, Castellano MA (1984) Una variante natural del virus della maculatura del garofano in Puglia. Inf Fitopatol 34:31–33 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Roggero P, Ciuffo M, Dellavalle G, Gotta P, Gallo S, Peters D (1999) Additional ornamental species as hosts of impatiens necrotic spot tospovirus in Italy. Plant Dis 83:967 Rubino L, Burgyan J, Russo M (1995) Molecular cloning and complete nucleotide sequence of carnation Italian ringspot tombusvirus genomic and defective interfering RNAs. Arch Virol 140(11):2027–2039 Ryabov EV, Generozov EV, Kendall TL, Lommel SA, Zavriev SK (1994) Nucleotide sequence of carnation ringspot dianthovirus RNA-1. J Gen Virol 75(1):243–247 Safari M, Koohi Habibi M, Mosahebi G, Dizadji A (2009) Carnation mottle virus, an important viral agent infecting carnation cut-flower crops in Mahallat of Iran. Commun Agric Appl Biol Sci 74(3):861–865 Sanchez-Navarro JA, Canizares MC, Cano EA, Pallas V (1999) Simultaneous detection of five carnation viruses by nonisotopic molecular hybridization. J Virol Methods 82:167–175 Sasaya T, Dujovny G, Koganezawa H (2000) Nucleotide sequence of the 30 – terminal region of carnation vein mottle virus RNA. J Gen Plant Pathol 66:251–253 Shahraeen N, Ghotbi T, Mehraban AH (2002) Occurrence of Impatiens necrotic spot virus in ornamentals in Mahallat and Tehran Provinces in Iran. Plant Dis 86:694 Singh HP, Hallan V, Raikhy G, Kulshrestha S, Sharma ML, Ram R, Garg ID, Zaidi AA (2005) Characterization of an Indian isolate of Carnation mottle virus infecting carnations. Curr Sci 88:594–601 Smookler M, Loebenstein G (1974) Carnation yellow fleck virus. Phytopathology 64:979–984 Stefanac Z, Wrischer M (1983) Occurrence of Carnation vein mottle and Cucumber mosaic viruses on carnations in Yugoslavia. Acta Bot Croat 42:21–27

D

880

Dianthus chinensis (China pink)

Szego A, Ilyes P, Toth EK, Potvondi L, Lukacs N (2006) Long term survival of cryptic viruses in aseptically grown in vitro propagated plants. Acta Hortic 725:505–510 Tremaine JH, Dodds JA (1985) Carnation ringspot virus. AAB descriptions of plant viruses, no. 308 Vainio EJ, Chiba S, Ghabrial SA, Maiss E, Roossinck M, Sabanadzovic S, Suzuki N, Xie J, Nibert M, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Partitiviridae. J Gen Virol 99:17–18 Wetter C (1971) Carnation latent virus. AAB descriptions of plant viruses, no. 61 White K (2011) Tombusvirus. Tombusviridae. In: The Springer index of viruses. Springer, New York, pp 1917–1926. https://doi.org/10.1007/978-0-387-95919-1_314 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354

Dianthus chinensis (China pink) Family: Caryophyllaceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Tospovirus

(INSV)

Family: Peribunyaviridae

INSV infection in plants of Dianthus chinensis was reported from Italy (Roggero et al. 1999). The virusinfected China pink plants show stunting, mosaic, and leaf malformation symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

References Roggero P, Ciuffo M, Dellavalle G, Gotta P, Gallo S, Peters D (1999) Additional ornamental species as hosts of impatiens necrotic spot tospovirus in Italy. Plant Dis 83:967

Diascia spp. Family: Scrophulariaceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Tospovirus

(INSV)

Family: Peribunyaviridae

INSV infection in plants of Diascia rigescens was reported from the USA (Daughtrey 1996). The virusinfected diascia plants exhibit necrotic patches and stem-streaking symptoms. The virus is transmitted

Diascia spp.

881

by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sapinoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Nemesia ring necrosis virus

(NeRNV)

Synonyms Diascia yellow mottle virus (DiaYMV)

Taxonomic position Genus: Tymovirus

D

Family: Tymoviridae

NeRNV infection was detected in plants of Diascia spp. in the USA, Germany, and the UK (Koenig et al. 2005; Mathews and Dodds 2006; Segwagwe et al. 2008; Harju et al. 2011). The virus-infected diascia plants exhibit symptoms ranging from non-symptomatic to a mild mosaic. The virus is transmitted by beetle vectors in a semi-persistent manner, and is also transmissible by mechanical sap-inoculation. Use of cuttings from infected plants for planting is the primary mode of spread. For more details of NeRNV, refer to Nemesia spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Tospovirus

(TSWV)

Family: Peribunyaviridae

TSWV infection in plants of Diascia barberae was reported by Parrella et al. (2003). The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Daughtrey ML (1996) Detection and identification of tospoviruses in greenhouses. Acta Hortic 431:90–98 Harju VA, Skelton AL, Monger WA, Forde SMD, Daly M, Nixon T, Lawson R, Bennett S, Mumford RA (2011) A report of viruses, viroids and phytoplasmas found in ornamental plants from 1999–2007: findings from Central Science Laboratory, UK. Acta Hortic 901:223–229 Koenig R, Barends S, Gultyaev AP, Lesemann DE, Vetten HJ, Loss S, Pleij CW (2005) Nemesia ring necrosis virus: a new tymovirus with a genomic RNA having a histidylatable tobamovirus-like 30 end. J Gen Virol 86:1827–1833 Mathews DM, Dodds JA (2006) First report of Nemesia ring necrosis virus in North America in ornamental plants from California. Plant Dis 90:1263 Parrella G, Gognalons P, Gebre-Selassi K, Vovlas C, Marchoux G (2003) An update of the host range of tomato spotted wilt virus. J Plant Pathol 85(4):227–264 Segwagwe AT, Putnam ML, Druffel KL, Pappu HR, Eastwell KC (2008) Molecular characterization of a new tymovirus from Diascia ornamental plants. Arch Virol 153:1495–1503

882

Dicentra spp. (Bleeding-heart)

Dicentra spp. (Bleeding-heart) Synonyms Lamprocapnos spectabilis Family: Papaveraceae

Ornamental

Arabis mosaic virus

(ArMV)

Taxonomic position Genus: Nepovirus

Family: Secoviridae

ArMV infection in plants of Dicentra formosa was reported from Lithuania (Samuitiene et al. 2008). The virus-infected bleeding heart plants show symptoms of plant chlorosis and stunting, with yellow ringspots on leaves. The virus is transmitted by the nematode vectors, Xiphinema diversicaudatum and X. coxi, in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in plants of Dicentra spectabilis was reported from Sweden and the USA (Lihnell and Nilsson 1969; Lockhart 2000; Fisher 2013). The virus-infected bleeding heart plants exhibit yellow ringspots and concentric line pattern symptoms. The virus is transmitted by nematode vectors and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Dicentra formosa was reported from Lithuania (Navalinskiene and Samuitiene 2000). The virus-infected bleeding heart plants show symptoms of chlorosis, stunting, and yellow ringspots on leaves. The virus is transmitted by nematode vectors in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and also by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Dichondra spp. (Dichondra)

883

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Tospovirus

(TSWV)

Family: Peribunyaviridae

TSWV infection in plants of Dicentra formosa was reported from Lithuania (Navalinskiene and Samuitiene 2006). The virus-infected bleeding heart plants show symptoms of chlorosis, stunting, and yellow ringspots on leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Fisher JR (2013) Identification of two Tobacco rattle virus variants associated with line pattern disease of bleeding heart in Ohio. Plant Health Prog. https://doi.org/10.1094/PHP-2013-0325-01-RS. Online Lihnell D, Nilsson B (1969) Vorkommen von Tabakrattle-virus an Dicentra spectabilis, in Schweden. J Phytopathol 65:1–6 Lockhart BEL (2000) Dicentra, Epimedium, and Heuchera: new perennial ornamental hosts of Tobacco rattle virus in the United States. Plant Dis 84:1344 Navalinskiene M, Samuitiene M (2000) Natural occurrence of tomato ringspot nepovirus in ornamental plants in Lithuania. In: Proceedings of the international conference: development of environmentally friendly plant protection in the Baltic Region, Tartu, Estonia, 28–29 Sept, pp 140–143 Navalinskiene M, Samuitiene M (2006) Dekoratyvinių augalų virusin_es ligos ir jų suk_el_ejai Lietuvoje. Lutut_e, Kaunas, 254 p Samuitiene M, Navalinskien_e M, Jackevičien_e E (2008) Arabis mosaic virus on ornamental plants. Biologija 54:264–268

Dichondra spp. (Dichondra) Family: Convolvulaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Weed host

(CMV)

Family: Bromoviridae

CMV infection in plants of Dichondra repens was reported from France, Italy, and China (Cardin et al. 2009). The virus-infected dichondra plants exhibit leaf mosaic and yellow ringspot symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of host plants. For more details of CMV, refer to Cucumis sativus.

D

884

Dichopogon strictus (Chocolate lily)

Turnip vein-clearing virus Taxonomic position Genus: Tobamovirus

(TVCV)

Family: Virgaviridae

TVCV infection in plants of Dichondra repens was reported from France, Italy, and China (Cardin et al. 2009). The virus-infected dichondra plants exhibit leaf mosaic and yellow ringspot symptoms. No insect vector is reported for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TVCV, refer to Brassica rapa.

References Cardin L, Delecolle B, Moury B (2009) First report of Cucumber mosaic virus and Turnip vein clearing virus in Dichondra repens in France, Italy and China. Plant Dis 93:201

Dichopogon strictus (Chocolate lily) Family: Asparagaceae

Chocolate lily virus A Taxonomic position Genus: Unassigned

Medicinal

(CLVA)

Family: Secoviridae

Geographical distribution CLVA infection in plants of Dichopogon strictus was reported from Australia (Wylie et al. 2012). Virion properties and genome The genome consists of two molecules of positive-sense single-stranded RNA. RNA1 consists of 6139 nt (JN052073 = NC_016443) and RNA2 of 4735 nt (JN052074 = NC_016444) (Wylie et al. 2012; Thompson et al. 2017).

References Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Secoviridae. J Gen Virol 98:529–531 Wylie SJ, Luo H, Li H, Jones MG (2012) Multiple polyadenylated RNA viruses detected in pooled cultivated and wild plant samples. Arch Virol 157(2):271–284

Dicliptera spp.

885

Dicliptera spp. Family: Acanthaceae

Weed host

Dicliptera yellow mottle Cuba virus Taxonomic position Genus: Begomovirus

(DiYMCUV)

D Family: Geminiviridae

Geographical distribution DiYMCUV infection in plants of Dicliptera vahliana was reported from Cuba (Echemendia et al. 2003). Symptoms and host(s) The virus-infected Dicliptera plants exhibit yellow mottle symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner, and also by grafting. The virus is not mechanically sap-transmissible. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2608 nt (AJ549960) (Echemendia et al. 2003; Zerbini et al. 2017).

Dicliptera yellow mottle virus Taxonomic Position Genus: Begomovirus

(DiYMoV)

Family: Geminiviridae

Geographical distribution DiYMoV infection in plants of Dicliptera spp. was reported from Cuba and Florida (Lotrakul et al. 2000; Echemendia et al. 2003). Symptoms and host(s) The virus-infected Dicliptera plants exhibit leaf distortion and yellow mottle symptoms. Transmission The virus has been shown to be transmitted by the whitefly vector of begomoviruses, Bemisia tabaci (Lotrakul et al. 2000). The mechanism of transmission has not been investigated. The virus is mechanically sap and graft-transmissible. Experimentally the virus has been shown to infect Nicotiana benthamiana, N. clevelandii, and D. stramonium (Lotrakul et al. 2000).

886

Dieffenbachia spp.

Virion properties and genome The structure of DiYMoV particles has not been investigated. In common with all geminiviruses, the virions of DiYMoV are likely geminate (twinned icosahedra). DiYMoV is typical of the majority of begomoviruses native to the New World with a bipartite genome consisting of two circular, single-stranded DNA components: DNA-A of 2607 nt (AF139168 = NC_003856) and DNA-B of 2597 nt (AF170101 = NC_003857) (Lotrakul et al. 2000; Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the New World, the DNA-A component of DiYMoV encodes five genes, one in the virionsense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for DiYMoV.

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Echemendia AL, Ramos PL, Peral R, Fuentes A, Pujol M, Gonzalez G (2003) First report of Dicliptera yellow mottle virus (DiYMoV) infecting Dicliptera vahliana in Cuba. Plant Pathol 52:787 Lotrakul P, Valverde RA, Landry AD (2000) Biological and molecular properties of a begomovirus from Dicliptera sexangularis. Phytopathology 90:723–729 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

Dieffenbachia spp. Family: Araceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

Family: Secoviridae

ArMV infection in plants of Dieffenbachia spp. was reported from Lithuania (Samuitiene et al. 2008). The virus-infected Dieffenbachia plants exhibit a slight leaf distortion and mosaic of irregular spots. The virus is transmitted by the nematode vectors Xiphinema diversicaudatum and X. coxi in a nonpersistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Clerodendrum chlorotic spot dichorhavirus Taxonomic position Genus: Dichorhavirus

Family: Rhabdoviridae

(ClCSV)

Dieffenbachia spp.

887

ClCSV infection in plants of Dieffenbachia amoena was reported from Brazil (Kitajima et al. 2008). The virus-infected Dieffenbachia plants exhibit chlorotic spot symptoms. The virus is transmitted by mite vectors (Brevipalpus spp.) and is also transmissible by mechanical sap-inoculation. For more details of ClCSV, refer to Clerodendrum spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

D Family: Bromoviridae

CMV infection in plants of Dieffenbachia spp. was reported from Czechoslovakia (Mokra and Gotzova 1994). The virus-infected Dieffenbachia plants exhibit symptoms of stunting, deformed leaves, and poor color clarity. The virus is transmitted by a number of aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of host plants. For more details of CMV, refer to Cucumis sativus.

Dasheen mosaic virus Taxonomic position Genus: Potyvirus

(DsMV)

Family: Potyviridae

DsMV infection in plants of Dieffenbachia spp. was reported from Australia, Europe, South Africa, South America, India, Brazil, and the USA (Hill and Wright 1980; Chase and Zettler 1982; Mokra and Gotzova 1994; Alexandre et al. 2005). In certain Dieffenbachia cultivars, DsMV induces inconspicuous symptoms causing little or no appreciable damage. In the case of D. maculata cultivars, symptoms range from slight mosaic to severe leaf distortion and also severe leaf size reduction (Elliott et al. 1997). The virus also induces prominent symptoms such as systemic mosaic, leaf malformation, vein-clearing, ringspots, and local necrosis. In some cultivars hypersensitivity is in the form of large spreading necrotic local lesions leads to the death of the plants. Symptoms of severity vary with time of year, species, and cultivar (Nelson 2008). The primary spread of DsMV takes place through use of propagating material from infected plants. Secondary spread takes place through aphid vectors in a non-spersistent manner, and is also transmissible through mechanical sap-inoculation (Zettler and Hartman 1987). For more details of DsMV, refer to Colocasia esculenta.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Tospovirus

(INSV)

Family: Peribunyaviridae

INSV infection in plants of Dieffenbachia amoena was reported from Iran (Ghotbi and Shahraeen 2012; Ghotbi 2013). The virus-infected Dieffenbachia plants exhibit symptoms of necrotic leaf spot, chlorosis, and stunting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

888

Konjac mosaic virus Taxonomic position Genus: Potyvirus

Dieffenbachia spp.

(KoMV)

Family: Potyviridae

KoMV infection in plants of Dieffenbachia spp. was reported from India and Taiwan (Padmavathi et al. 2011). The virus-infected Dieffenbachia plants exhibit symptoms of mosaic, chlorotic feathery mottling, chlorotic spots, leaf deformation, and chlorotic ringspots. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of KoMV, refer to Amorphophallus paeoniifolius.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV infection in plants of Dieffenbachia spp. was reported from Brazil and Czechoslovakia (Mokra and Gotzova 1994; Rivas et al. 2000). The virus-infected Dieffenbachia plants exhibit chlorotic spots and rings and leaf deformation symptoms. There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tomato chlorotic spot orthotospovirus Taxonomic position Genus: Tospovirus

(TCSV)

Family: Peribunyaviridae

TCSV infection in plants of Dieffenbachia amoena was reported from Brazil (Rivas et al. 2002). The virus-infected Dieffenbachia plants exhibit symptoms of vein-banding, necrotic lines, and chlorotic spots and rings. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of TCSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Tospovirus

(TSWV)

Family: Peribunyaviridae

TSWV was reported in plants of Dieffenbachia amoena in Poland and Iran (Kaminska and Korbin 1994; Ghotbi and Shahraeen 2012). The virus-infected Dieffenbachia plants show chlorotic and/or necrotic spots and rings or line patterns. The virus is transmitted by thrips vectors in a persistentpropagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Digitalis spp. (Foxglove)

889

Tomato yellow ring virus

(TYRV)

Taxonomic position TYRV is a tentative member of the genus Tospovirus and family Peribunyaviridae TYRV was detected in plants of Dieffenbachia amoena in Iran (Ghotbi and Shahraeen 2012). The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of TYRV, refer to Solanum lycopersicum.

References Alexandre MAV, Rivas EB, Tozetto ARP, Duarte LML (2005) An annotated list on the natural occurrence of viruses in ornamental plants in Brazil. Instituto Biologico, Sao Paulo, 54 p Chase AR, Zettler FW (1982) Dasheen mosaic virus infection of dieffenbachia cultivars. Plant Dis 66:891–893 Elliott MS, Zettler FW, Brown LG (1997) Dasheen mosaic potyvirus of edible and ornamental aroids. Plant pathology circular No. 384, Fla Dept and Consumer Services, July/August 1997 Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iran J Plant Pathol 49:41 Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses affecting propagated ornamentals in Northern Iran. Int Res J Microbiol 3:373–381 Hill SA, Wright DM (1980) Identification of Dasheen mosaic virus in Dieffenbachia picta and Xanthosoma helliborifolium by immune electron microscopy. Plant Pathol 29:143–144 Kaminska M, Korbin M (1994) New natural hosts of Tomato spotted wilt virus. Acta Hortic 377:123–128 Kitajima EW, Kubo KS, Ferreira PTO, Alc^antara BK, Boari AJ, Gomes RT, Freitas-Astúa J, Rezende JAM, Morais GJ, Salaroli RB (2008) Chlorotic spots on Clerodendrum, a disease caused by a nuclear type of Brevipalpus (Acari: Tenuipalpidae) transmitted virus. Sci Agric 65:36–49 Mokra V, Gotzova B (1994) Identification of virus infections in Dieffenbachia and Zantedeschia in Czechoslovakia. Acta Hortic 377:361–362 Nelson SC (2008) Dasheen mosaic of edible and ornamental aroids. Plant disease CTAHR. Available at: http://www.ctahr. hawaii.edu/oc/freepubs/pdf/PD-44.pdf Padmavathi M, Srinivas KP, Subba Reddy CV, Ramesh B, Navodayam K, Krishnaprasadji J, Babu Ratan P, Sreenivasulu P (2011) Konjac mosaic virus naturally infecting three aroid plant species in Andhra Pradesh, India. J Phytopathol 159:133–135 Rivas EB, Pezani EF, Alexandre MAV, Duarte LML (2000) First report of a Tobamovirus in dieffenbachia and impatiens. Plant Dis 84:707 Rivas EB, Galleti SR, Alexandre MAV, Duarte LML, Seabra PV, Estelita MEM (2002) Tomato chlorotic spot virus (Tospovirus) in dieffenbachia and bouvardia. Virus Rev Res 7:22–30 Samuitiene M, Navalinskiene M, Jackeviciene E (2008) Arabis mosaic virus on ornamental plants. Biologia 54:264–268 Zettler FW, Hartman RD (1987) Dasheen mosaic virus as a pathogen of cultivated aroids and control of the virus by tissue culture. Plant Dis 71:958–963

Digitalis spp. (Foxglove) Family: Plantaginaceae

Broad bean wilt virus 2 Taxonomic position Genus: Fabavirus

Medicinal

(BBWV-2)

Family: Secoviridae

D

890

Digitalis spp. (Foxglove)

BBWV-2 infection in plants of Digitalis spp. was reported from the United Kingdom, Germany, China, and Yugoslavia (Schumann 1963a; Milicic et al. 1976; Mumford et al. 2006; Dong et al. 2017). The virus-infected foxglove plants show mottling symptoms on the leaves. The virus is transmitted by an aphid vector Acyrthosiphon solani in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BBWV-2, refer to Vicia faba.

Ribgrass mosaic virus

(RMV)

Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

RMV infection in plants of Digitalis spp. was reported from Germany and Yugoslavia (Schumann 1963b; Juretic and Milicic 1977). The virus-infected foxglove plants exhibit mosaic symptoms. No vector is reported for this virus. The virus is mechanically sap-transmissible. The virus is transmissible by grafting and also by contact between plants. For more details of RMV, refer to Plantago lanceolata.

Tobacco mosaic virus

(TMV)

Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

TMV infection in plants of Digitalis thapsi was reported from Spain (Vela 1972). There is no known vector for this virus. The virus is mechanically sap-transmissible and is also transmissible through contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tomato bushy stunt virus Taxonomic position Genus: Tombusvirus

(TBSV)

Family: Tombusviridae

TBSV infection in plants of Digitalis grandiflora was reported by Novak and Lanzova (1980). The virus is transmissible by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants. There is no known vector for this virus. For more details of TBSV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Digitalis purpurea was reported from Lithuania (Navalinskiene and Samuitiene 2000). The virus is transmitted by nematode vectors in a non-persistent manner, and is also

Digitalis spp. (Foxglove)

891

transmissible by mechanical sap-inoculation, and by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Tospovirus

(TSWV)

Family: Peribunyaviridae

TSWV infection in plants of Digitalis purpurea was reported from Pennsylvania (USA) (Hausbeck et al. 1992). The virus-infected foxglove plants exhibit mottle mosaic, necrosis, and chlorotic spotting symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Youcai mosaic virus Taxonomic position Genus: Tobamovirus

(YoMV)

Family: Virgaviridae

YoMV infection in plants of Digitalis lanata was reported from Japan (Dong et al. 2017). There is no known vector for this virus. The virus is mechanically sap-transmissible and is also transmissible through contact between plants. For more details of YoMV, refer to Brassica napus.

References Dong JL, Li Y, Ding WL, Wang R (2017) First report of broad bean wilt virus 2 and Youcai mosaic virus infecting woolly foxglove (Digitalis lanata). J Plant Pathol 99:816 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among green house ornamentals in Pennsylvania. Plant Dis 76:795–800 Juretic N, Milicic D (1977) Natural infection of Digitalis ciliata with ribgrass mosaic virus. Acta Bot Croat 36:23–28 Milicic D, Juretic N, Plese N, Wrischer M (1976) Some data on cell inclusions and natural hosts of broad bean wilt virus. Acta Bot Croat 35:17 Mumford RA, Jarvis B, Harju V, Boonham N, Skelton A (2006) First report of Broad bean wilt virus – 2 in the UK: Findings in foxglove and salvia. Plant Pathol 55:819 Navalinskiene M, Samuitiene M (2000) Natural occurrence of tomato ringspot nepovirus in ornamental plants in Lithuania. In: Proceedings of the international conference: development of environmentally friendly plant protection in the Baltic Region, Tartu, Estonia, 28–29 Sept, pp 140–143 Novak JB, Lanzova J (1980) Demonstration of tomato bushy stunt virus in some forest tree species and plants. Lesnictví 26:1009–1016 Schumann K (1963a) Untersuchungen zur Charakterisierung und Identifizierung der Erreger des Digitalis-Mosaik. II. Das Ringmosaik-Virus der Kapuzinerkresse. Phytopathol Z 48:135–148 Schumann K (1963b) Untersuchungen zur Charakterisierung and Identifizierung der Erreger des “Digitalis-Mosaik.” I. Das Tabakmosaikvirus (Marmor tabaci H.). Phytopathol Z 48:1–28 Vela A (1972) Study of a strain of tobacco mosaic virus from digitalis thapsi L. Microbiol Espa 25:211–224

D

892

Digitaria spp. (Pangola grass)

Digitaria spp. (Pangola grass) Family: Poaceae

Forage crop

Barley yellow dwarf virus PAV Taxonomic position Genus: Luteovirus

(BYDV-PAV)

Family: Luteoviridae

BYDV-PAV infection in plants of Digitaria sanguinalis was reported from Western Australia (Jones et al. 1990). The virus is transmitted by a number of aphid species in a circulative, non-propagative manner. No mechanical transmission of this virus is reported. For more details of BYDV-PAV, refer to Hordeum vulgare.

Cereal yellow dwarf virus RPV Taxonomic position Family: Luteoviridae

(CYDV-RPV)

Genus: Polerovirus

CYDV-RPV infection in Digitaria sanguinalis plants was reported from Western Australia (Jones et al. 1990). The virus is transmitted by a number of aphid species in a circulative, non-propagative manner. No mechanical transmission of this virus is reported. For more details of CYDV-RPV, refer to Hordeum vulgare.

Digitaria ciliaris striate mosaic virus Taxonomic position Genus: Mastrevirus

(DCSMV)

Family: Geminiviridae

Geographical distribution DCSMV infection in plants of Digitaria spp. was reported from Australia and Africa (Kraberger et al. 2012). Symptoms and host(s) The virus-infected pangola grass plants exhibit striate mosaic symptoms. Transmission The virus is transmitted by leafhopper vectors in a persistent, circulative, and non-propagative manner. The virus is not transmissible by mechanical inoculation. Virion properties and genome The virions are twinned (geminated) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is a monopartite circular single-stranded DNA of 2816 nt (JQ948088 = NC_018579) (Palmer and Rybicki 1998; Boulton and Davies 2011; Kraberger et al. 2012; Muhire et al. 2013; Zerbini et al. 2017).

Digitaria spp. (Pangola grass)

893

Digitaria didactyla striate mosaic virus Taxonomic position Genus: Mastrevirus

(DDSMV)

Family: Geminiviridae

Geographical distribution DDSMV was first reported in plants of Digitaria didactyla from Queensland, Australia, by Greber (1989). The virus spreads in Australia (Briddon et al. 2010). Symptoms and host(s) The virus-infected pangola grass plants exhibit fine striations and a blotchy mottle which may cover the whole leaf surface. Transmission The virus is transmitted by the leafhopper Nesoclutha pallida (Grylls 1979). In common with all other geminiviruses, DDSMV is likely transmitted in a circulative, non-propagative manner. The virus is not transmissible by mechanical inoculation (Greber 1989). Experimentally the virus was transferred to Aegilops variabilis, Avena sativa, Bromus catharticus, Dactyloctenium aegyptium, Hordeum vulgare, Leptochloa filiformis, and Triticum aestivum (Greber 1989). Virion properties and genome Purified virus DDSMV preparations contained virions that are twinned icosahedra, typical of geminiviruses (Pinner et al. 1992). The genome is a monopartite circular single-stranded DNA of 2762 nt (HM122238 = NC_014547) (Pinner et al. 1992; Palmer and Rybicki 1998; Briddon et al. 2010; Boulton and Davies 2011; Muhire et al. 2013; Zerbini et al. 2017). In common with all mastreviruses, DDSMV is predicted to encode four gene products, two in each orientation. Based on sequence homology to previously studied mastreviruses, these would code for the coat protein and movement protein in the virion-sense and in the complementary-sense for the replication-associated protein (Rep) and Rep A.

Digitaria streak virus Taxonomic position Genus: Mastrevirus

(DSV)

Family: Geminiviridae

Geographical distribution DSV infection was first reported in plants of Digitaria setigera (Syn.) Digitaria sanguinalis from Vanuatu by Dollet et al. (1986). The virus spreads in Vanuatu (Donson et al. 1987; Julia and Dollet 1989). Symptoms and host(s) The virus-infected pangola grass plants exhibit narrow white streaks and chlorotic local lesions with diffuse margins.

D

894

Digitaria spp. (Pangola grass)

Transmission The virus is transmitted by the leafhopper vector Nesoclutha declivata in a circulative and nonpropagative manner (Julia and Dollet 1989). The virus is not transmissible by mechanical inoculation, grafting, or contact between plants. The virus is not transmitted by seed and pollen. Experimentally the virus was transmissible to D. setigera by insect transmission, as well Zea mays and Avena sativa by Agrobacterium-mediated inoculation of cloned viral DNA (Donson et al. 1988). Virion properties and genome Purified virus preparations of DSV contained virions that were twinned icosahedra (geminated), typical of geminiviruses, with a size of approx. 20  30 nm (Dollet et al. 1986). Only a single isolate of DSV has been characterized (Donson et al. 1987). The genome consists of single circular single-stranded DNA of 2701 nt (M23022 = NC_001478) (Donson et al. 1987, 1988; Accotto et al. 1989; Palmer and Rybicki 1998; Boulton and Davies 2011; Muhire et al. 2013; Zerbini et al. 2017). In common with all mastreviruses, DSV is predicted to encode four gene products, two in each orientation. Based on sequence homology to previously studied mastreviruses, these would code for the coat protein and movement protein in the virion-sense and in the complementary-sense for the replication-associated protein (Rep) and Rep A. The transcription of the DSV genome occurs in both orientations (bidirectional) and one of the transcripts covering the complementary-sense is spliced, allowing the production of two products (Rep from a spliced transcript and Rep A from a non-spliced transcript) (Accotto et al. 1989).

Digitaria striate virus

(DiSV)

Taxonomic position DiSV is a tentative member of the family Rhabdoviridae Geographical distribution DiSV infection was first reported in plants of Digitaria decumbens and D. ciliaris from the coastal areas of northern and southern Queensland by Greber (1972). The virus spreads in Australia (Greber 1979, 1996). Symptoms and host(s) The virus-infected pangola grass exhibit symptoms of striate and chlorotic spots and stripes. The natural host range of this virus is Urochloa distachya, Digitaria decumbens, D. ciliaris, D. eriantha, D. sanguinalis, and Eleusine indica. Transmission The virus is transmitted by a planthopper vector Sogatella kolophon in a persistent and propagative manner. Virus is retained through the vector molts. The virus is acquired within a 24-h acquisition feeding period, and the latent period in the insect is from 5 to 8 days. The virus is not transmissible by mechanical inoculation. The virus is not transmitted by seed or pollen. Virion properties and genome The virions are bullet shaped, enveloped with a clear modal length of 280 nm and 55 nm wide. The genome is a monopartite, negative-sense, single-stranded RNA (Greber 1990).

Digitaria spp. (Pangola grass)

895

Pangola stunt virus Taxonomic position Genus: Fijivirus

(PaSV)

Family: Reoviridae

Geographical distribution PaSV was first reported in plants of Digitaria decumbens from Surinam by Dirven and van Hoof (1960). The virus spreads in Australia, Brazil, Fiji, Guyana, Peru, and Taiwan (Teakle et al. 1988, 1991). Symptoms and host(s) The virus-infected pangola grass plants exhibit symptoms of stunting, distortion of leaves and inflorescences, and poor regeneration after cropping. Transmission The virus is transmitted by a planthopper vectors Sogatella furcifera and S. kolophon in a propagative manner. The shortest acquisition time is 30–60 min, the latent period is 3–4 weeks, and the vectors remain viruliferous throughout life. No transovarial or seed-transmission of the virus has been identified. The virus is not transmissible by mechanical inoculation, by grafting, by contact between plants, or by pollen (Teakle et al. 1988). Virion properties and genome The virions are non-enveloped and icosahedral with a double-capsid structure, about 65–70 nm in diameter. Pentameric turrets sit on the outside of the innermost capsid. The outer capsid has a T = 13 icosahedral symmetry, the inner capsid a T = 2 icosahedral symmetry. The genome is a linear doublestranded RNA with 10 segments coding for 12 proteins (Francki and Boccardo 1983; Karan et al. 1994; Marzachi et al. 1995; Harding and Dale 2011).

Sugarcane mosaic virus Taxonomic position Genus: Potyvirus

(SCMV)

Family: Potyviridae

SCMV infection in plants of Digitaria nuda and D. abyssinica was reported from Kenya (Louie 1980). The virus-infected pangola grass plants exhibit chlorotic streaking symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of SCMV, refer to Saccharum officinarum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Tospovirus

(TSWV)

Family: Peribunyaviridae

TSWV infection in plants of Digitaria sanguinalis was reported from Italy (Grieco et al. 2000). The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

D

896

Digitaria spp. (Pangola grass)

References Accotto G-P, Donson J, Mullineaux PM (1989) Mapping of Digitaria streak virus transcripts reveals different RNA species from the same transcription unit. EMBO J 8:1033–1039 Boulton MI, Davies JW (2011) Mastrevirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 597–603. https://doi.org/10.1007/978-0-387-95919-1_83 Briddon RW, Martin DP, Owor BE, Donaldson L, Markham PG, Greber RS, Varsani A (2010) A novel species of mastrevirus (family Geminiviridae) isolated from Digitaria didactyla grass from Australia. Arch Virol 155:1529–1534 Dirven JGP, Van Hoof HA (1960) A destructive virus disease of pangola-grass. Tijdschrift Over Plantenziekten 66:344–349 Dollet M, Accotto GP, Lisa V, Menissier J, Boccardo G (1986) A geminivirus serologically related to maize streak virus, from Digitaria sanguinalis from Vanuatu. J Gen Virol 67:933–937 Donson J, Accotto GP, Boulton MI, Mullineaux PM, Davies JW (1987) The nucleotide sequence of a geminivirus from Digitaria sanguinalis. Virology 161:160–169 Donson J, Gunn HV, Woolston CJ, Pinner MS, Boulton MI, Mullineaux PM, Davies JW (1988) Agrobacterium-mediated infectivity of cloned Digitaria streak virus DNA. Virology 162:248–250 Francki RIB, Boccardo G (1983) The plant reoviridae. In: Joklik WK (ed) The reoviridae. Plenum Press, New York, pp 505–563 Greber RS (1972) Particles of the rhabdovirus type in striate diseased pangola grass. Aust Plant Pathol Soc Newsl 1:27–28 Greber RS (1979) Digitaria striate virus a rhabdovirus of grasses transmitted by Sogatella kolophon. Aust J Agric Res 30:43–51 Greber RS (1989) Biological characteristics of grass geminiviruses from eastern Australia. Ann Appl Biol 114:471–480 Greber RS (1990) Discrimination by gel-diffusion serology of Digitaria striate, maize sterile stunt and other rhabdoviruses of Poaceae. Ann Appl Biol 116:259–264 Greber RS (1996) Digitaria striate cytorhabdovirus. In: Brunt AA et al (eds) Viruses of plants. CAB International, Wallingford, pp 538–539 Grieco PD, Conte D, Munno I, Nuzzaci M, de Stradis A (2000) Tomato spotted wilt virus (TSWV) on weeds and wild plants in Metaponto, Basilicata. Informatore Fitopatologico 50:43–46 Grylls NE (1979) Leafhopper vectors and the plant disease agents they transmit in Australia. In: Maramorosch K, Harris KF (eds) Leafhopper vectors and plant disease agents, 0th edn. Academic, New York, pp 179–214 Harding RM, Dale JL (2011) Fijivirus. Reoviridae. In: The Springer index of viruses. Springer, New York, pp 1589–1593. https://doi.org/10.1007/978-0-387-95919-1_260 Jones RAC, McKirdy SJ, Shivas RG (1990) Occurrence of barley yellow dwarf viruses in over-summering grasses and cereal crops in Western Australia. Aust Plant Pathol 19:90–96 Julia JF, Dollet M (1989) Nesoclutha declivata Homoptera Cicadellidae, vector of Digitaria streak virus (Geminivirus) in Vanuatu. J Phytopathol 127:42–48 Karan M, Dale JL, Bateson MF, Harding RM, Teakle DS (1994) Detection and characterization of pangola stunt Fijivirus from Australia using cloned cDNA probes. Arch Virol 135:397–404 Kraberger S, Thomas JE, Geering AD, Dayaram A, Stainton D, Hadfield J, Walters M, Parmenter KS, van Brunschot S, Collings DA, Martin DP, Varsani A (2012) Australian monocot-infecting mastrevirus diversity rivals that in Africa. Virus Res 169(1):127–136 Louie R (1980) Sugarcane mosaic virus in Kenya. Plant Dis 64:944–947 Marzachi C, Boccardo G, Milne R, Isogai M, Uyeda I (1995) Genome structure and variability of fijiviruses. Semin Virol 6:103–108 Muhire B, Martin DP, Brown JK, Navas-Castillo J, Moriones E, Zerbini FM, Rivera-Bustamante R, Malathi VG, Briddon RW, Varsani A (2013) A genome-wide pairwise-identity-based proposal for the classification of viruses in the genus Mastrevirus (family Geminiviridae). Arch Virol 158(6):1411–1424 Palmer KE, Rybicki EP (1998) The molecular biology of mastreviruses. Adv Virus Res 50:183–234 Pinner MS, Markham PG, Rybicki EP, Greber RS (1992) Serological relationships of Geminivirus isolates from Gramineae in Australia. Plant Pathol 41:618–625 Teakle DS, Hicks S, Harding RM, Greber RS, Milne RG (1988) Pangola stunt virus infecting pangola grass and summer grass in Australia. Aust J Agric Res 39:1075–1083 Teakle DS, Hicks S, Karan M, Hacker JB, Greber RS, Donaldson JF (1991) Host range and geographic distribution of pangola stunt virus and its planthopper vectors in Australia. Austral J Agric Res 42(5):819–826 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

Dimorphotheca sinuata (Glandular Cape marigold)

897

Dimocarpus longan (Longan) Family: Sapindaceae

Edible fruit

Longan witches broom-associated virus (LWBDV) Taxonomic position Genus: Unassigned Family: Potyviridae Geographical distribution LWBDV infection in plants of Dimocarpus longan was reported from Hanoi, Vietnam (Seo et al. 2017). Symptoms and host(s) The virus-infected longan plants show witches' broom syndrome. Transmission There is no known vector for this virus. The virus is transmissible by mechanical sap-inoculation. Virion properties and genome The virions are flexuous filaments, with an average length of 730 nm and 12 nm wide. The genome consists of linear positive-sense single-stranded RNA of 9428 nt (KY649478 = NC_034835) excluding the 30 poly(A) tail and contains one large single open reading frame encoding a polyprotein of 3086 amino acids (Seo et al. 2017; Wylie et al. 2017).

References Seo JK, Kim MK, Kwak HR, Kim JS, Choi HS (2017) Complete genome sequence of longan witches’ broom-associated virus, a novel member of the family potyviridae. Arch Virol 162(9):2885–2889 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: potyviridae. J Gen Virol 98:352–354

Dimorphotheca sinuata (Glandular Cape marigold) Family: Asteraceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Ornamental

(AMV)

Family: Bromoviridae

D

898

Dimorphotheca sinuata (Glandular Cape marigold)

AMV was reported infecting plants of Dimorphotheca sinuata in Germany (Kleinhempel 1991). The virus is transmitted by a large number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Dimorphotheca sinuata was reported from Italy (Dellavalle et al. 1994). The virus-infected glandular Cape marigold plants show distorted leaves with green mosaic. The infected plants were stunted and yielded fewer flowers. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Lettuce mosaic virus Taxonomic position Genus: Potyvirus

(LMV)

Family: Potyviridae

LMV was reported in plants of Dimorphotheca sinuata from Greece, Japan, and Italy (Lisa 1995; Manoussopoulos et al. 1999). The virus-infected glandular Cape marigold plants show symptoms of numerous yellow patches appearing mainly on the lower leaves and yellow rings on the upper and nascent leaves. Growth of the diseased plants was slightly retarded, but they flowered normally. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of LMV, refer to Lactuca sativa.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Tospovirus

(TSWV)

Family: Peribunyaviridae

TSWV infection in plants of Dimorphotheca sinuata was reported from Greece, France, Italy, and Japan (Ramasso et al. 1994: Lisa 1995; Manoussopoulos et al. 1999; Uga and Tsuda 2005). The virusinfected glandular Cape marigold plants show generalized yellowing, necrosis, and mosaic on leaves and sometimes tip necrosis. In early infected plants, many are killed, while plants infected at later stages can survive but are unmarketable. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Diodia virginiana (Virginia buttonweed)

899

References Dellavalle G, Masenga V, Lisa V (1994) Cucumber mosaic, a Potyvirus and a Carlavirus from Dimorphotheca sinuata. Acta Hortic 377:91–98 Kleinhempel K (1991) Virus infection of outdoor ornamental plants. In: Rep. for 1990, Inst. für Phytopathologie, Aschersleben, p 36 Lisa V (1995) Dimorphotheca sinuata. In: Loebenstein G, Lawson RH, Brunt AA (eds) Virus and virus-like diseases of bulb and flower crops. Wiley, New York, pp 471–475 Manoussopoulos IN, Chatzivassiliou EK, Smyrnioudis IN, Katis NI (1999) Two diseases of Dimorphotheca caused by lettuce mosaic potyvirus and tomato spotted wilt tospovirus. Phytoparasitica 27:227–232 Ramasso E, Dellavalle G, Roggero P, Lisa V (1994) Nuovi ospiti di Tospovirus in piante ornamentali ed ortensi in Liguria. Inf Fitopatol 44:44–48 Uga H, Tsuda S (2005) A one-step reverse transcription-polymerase chain reaction system for the simultaneous detection and identification of multiple tospovirus infections. Phytopathology 95:166–171

Diodia virginiana (Virginia buttonweed) Family: Rubiaceae

Weed

Diodia vein chlorosis virus

(DVCV)

Synonyms Diodia yellow vein virus Taxonomic position Genus: Crinivirus

Family: Closteroviridae

Geographical distribution DVCV was first identified by Larsen et al. (1991) in Diodia virginiana. DVCV (also known as Diodia yellow vein virus) is present in the USA and Australia (Tzanetakis et al. 2011, 2013). Symptoms and host(s) The virus-infected Virginia buttonweed plants exhibit chlorosis and vein-netting symptoms (Larsen et al. 1991). The natural host range of this virus is Diodia virginiana. Transmission The virus is transmitted by the whitefly vectors Trialeurodes abutilonea and T. vaporariorum in a semipersistent manner; they transmit the virus with efficiencies of over 36% and 12%, respectively, when plants were inoculated with 50 whiteflies after 48-h AAP and IAP (Tzanetakis et al. 2011). The virus is not mechanically sap-transmissible but transmissible by grafting. Virion properties and genome The virions are non-enveloped, bipartite filaments particles about 650–850 nm and 700–900 nm in length, and 10–13 nm in diameter. The genome consists of two molecules of positive-sense ssRNA. RNA-1 consists of 8010 nt (GQ225585) and RNA-2 of 8220 nt (GQ376201) (Kreuze 2011; Tzanetakis et al. 2011).

D

900

Dioscorea spp. (Yam)

References Kreuze JF (2011) Crinivirus. Closteroviridae. In: The Springer index of viruses. Springer, New York, pp 335–342. https:// doi.org/10.1007/978-0-387-95919-1_51 Larsen RC, Kim KS, Scott HA (1991) Properties and cytopathology of Diodia vein chlorosis virus – a new whiteflytransmitted virus. Phytopathology 81:227–232 Tzanetakis IE, Wintermantel WM, Poudel B, Zhou J (2011) Diodia vein chlorosis virus is a group-1 crinivirus. Arch Virol 156:2033–2037 Tzanetakis IE, Martin RR, Wintermantel WM (2013) Epidemiology of Criniviruses: an emerging problem in world agriculture. Front Microbiol 4:119. https://doi.org/10.3389/fmicb.2013.00119

Dioscorea spp. (Yam) Family: Dioscoreaceae

Tuber crop

Air potato ampelovirus 1 Taxonomic position Genus: Ampelovirus

(AiPoV-1)

Family: Closteroviridae

Geographical distribution AiPoV-1 infection in plants of air potato (Dioscorea bulbifera) was reported from the USA (Dey et al. 2019). Symptoms and host range The virus-infected plants exhibit foliar mosaic symptoms. Transmission The virus is transmitted by mealybug vectors. The virus is not transmitted by mechanical sapinoculation, and through seed. The virus is transmissible by grafting. Virion properties and genome The virions are filamentous, non-enveloped, very flexuous, 1500 nm long and 10–13 nm in diameter. The genome is a monopartite, linear, single-stranded, positive-sense RNA of 13398 nt (MH206615) possessing seven open reading frames (ORFs) (Dey et al. 2019).

Broad bean wilt virus 2 Taxonomic position Genus: Fabavirus

(BBWV-2)

Family: Secoviridae

BBWV-2 infection in plants of Dioscorea opposita was reported from Japan and Korea (Ishikawa et al. 1985; Kondo et al. 2005; Kwon et al. 2016). The virus-infected yam plants exhibit symptoms of veinclearing followed by mosaic and rugose symptoms. The outer parts of the leaf roll down towards the under-side and are malformed. The virus is transmitted by an aphid vector Acyrthosiphon solani in a

Dioscorea spp. (Yam)

901

non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BBWV-2, refer to Vicia faba.

Chinese yam necrotic mosaic virus Taxonomic position Genus: Macluravirus

(CYNMV)

Family: Potyviridae

Geographical distribution CYNMV infection in plants of Dioscorea opposita cv. nagaimo was reported from Japan (Fukumoto and Tochihara 1978). This virus is quite prevalent in China and Japan (Wang et al. 2009; Kondo and Fujita 2012).

Symptoms and host(s) The virus-infected yam plants exhibit symptoms of conspicuous chlorotic and/or necrotic spots, mosaic and netting, and severe interveinal chlorosis and necrosis. The virus is restricted to Chinese yam, and does not infect any other host.

Transmission The virus is transmitted efficiently in the non-persistent manner by aphids (especially Aphis gossypii and Myzus persicae). The virus is also mechanically sap-transmissible.

Virion properties and genome The virions are non-enveloped, flexuous filaments, 660 nm long, and 13–16 nm in diameter, with a coat protein of 38 kDa. The genome consists of a single molecule of linear positive-sense ssRNA of 8224 nt (AB710145 = NC_018455) (Kondo 2001; Kondo et al. 2003; Foster 2011; Kondo and Fujita 2012; Wylie et al. 2017).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Dioscorea alata was reported from West Africa (Fauquet and Thouvenel 1987). Strains of CMV have been reported infecting D. alata, D. trifida, and D. rotundata in West Africa, the Caribbean, Ghana, Nigeria, Togo, South America, and South Pacific (Migliori and Cadilhac 1976; Hughes et al. 1997; Eni et al. 2008a, c, 2010, 2013; Odedara et al. 2011; Asala et al. 2012). The virus-infected yam plants exhibit severe leaf chlorosis and mosaic symptoms. The virus also causes leaf distortions and stunting. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a wide range of indicator hosts. For more details of CMV, refer to Cucumis sativus.

D

902

Dioscorea spp. (Yam)

Dioscorea bacilliform AL virus Synonyms Dioscorea alata bacilliform virus (DaBV); Dioscorea bacilliform virus; Dioscorea bulbifera virus (DBALV); Yam internal brown spot virus Taxonomic position Genus: Badnavirus

Family: Caulimoviridae

Geographical distribution DBALV infection in plants of Dioscorea spp. was reported from the Caribbean area, Barbados, West Africa, Asia, Nigeria, and South American countries (Harrison and Roberts 1973; Seal and Muller 2007; Eni et al. 2008c; Odedara et al. 2011). Symptoms and host(s) The virus-infected yam plants exhibit symptoms of severe interveinal leaf chlorosis in D. bulbifera and leaf crinkling, distortion, vein chlorosis, and vein necrosis, associated with internal brown spotting of tubers in D. alata cv. White Lisbon. Transmission The virus is transmitted by the mealybug vector Planococcus citri in a semi-persistent manner. The virus is transmissible by mechanical sap-inoculation of partially purified preparations to several other Dioscorea species, whereas inoculation to herbaceous hosts outside the Dioscoreaceae failed. Use of infected vegetative propagative material is another source of virus spread. Virion properties and genome The virions are bacilliform, non-enveloped, and about 28 nm in diameter and 130 nm in length. The genome is a monopartite, circular double-stranded DNA of 7.5 kbp (X94576) with a single-stranded discontinuity at one site in each strand. The coat protein of DaBV was about 56 kDa (Briddon et al. 1999; Phillips et al. 1999; Olszewski and Lockhart 2011; Bhat et al. 2016).

Dioscorea bacilliform AL virus 2 Taxonomic position Genus: Badnavirus

(DBALV2)

Family: Caulimoviridae

Geographical distribution DBALV2 infection in plants of Dioscorea alata was reported from Papua New Guinea (Sukal et al. 2017). Transmission The virus is transmitted by mealybug vectors in a semipersistent manner. The virus is transmitted by mechanical inoculation (with difficulty). The virus is transmitted by grafting and not by contact between plants or by pollen.

Dioscorea spp. (Yam)

903

Virion properties and genome The virions are bacilliform in shape with a diameter of 30 nm and modal particle length of 130 nm. The genome is a circular dsDNA of 7871 bp (KY827395) with three open reading frames (ORFs) (Olszewski and Lockhart 2011; Bhat et al. 2016; Sukal et al. 2017).

Dioscorea bacilliform ES virus Taxonomic position Genus: Badnavirus

(DBESV)

D Family: Caulimoviridae

Geographical distribution DBESV infection in plants of Dioscorea esculenta was reported from Fiji (Sukal et al. 2017). Transmission The virus is transmitted by mealybug vectors in a semipersistent manner. The virus is transmitted by mechanical inoculation (with difficulty). The virus is transmitted by grafting and not by contact between plants or by pollen. Virion properties and genome The virions are bacilliform in shape with a diameter of 30 nm and modal particle length of 130 nm. The genome is a circular dsDNA of 8106 bp (KY827394) with three open reading frames (ORFs) (Olszewski and Lockhart 2011; Bhat et al. 2016; Sukal et al. 2017).

Dioscorea bacilliform RT virus 1 Taxonomic position Genus: Badnavirus

(DBRTV1)

Family: Caulimoviridae

Geographical distribution DBRTV1 infection in plants of Dioscorea rotundata was reported from Nigeria (Bomer et al. 2016). Transmission The virus-infected planting material is the main source of virus spread. The virus is not mechanically sap-transmissible. Virion properties and genome Virions are bacilliform in shape with a diameter of 30 nm and modal particle length of 130 nm (Olszewski and Lockhart 2011). The genome contains a single molecule of non-covalently closed circular dsDNA of 7702 bp (KX008574 = NC_038986) with 3 ORFs, including a large ORF (ORF3) encoding a putative polyprotein with a conserved movement protein, coat protein, aspartic protease, reverse transcriptase, and ribonuclease H domains analogous to those of other badnaviruses (Bomer et al. 2016).

904

Dioscorea spp. (Yam)

Dioscorea bacilliform RT virus 2 Taxonomic position Genus: Badnavirus

(DBRTV2)

Family: Caulimoviridae

Geographical distribution DBRTV2 infection in plants of Dioscorea rotundata was reported from Nigeria (Bomer et al. 2016). Transmission The virus-infected planting material is the main source of virus spread. The virus is not mechanically sap-transmissible. Virion properties and genome Virions are bacilliform in shape with a diameter of 30 nm and modal particle length of 130 nm (Olszewski and Lockhart 2011). The genome contains a single molecule of non-covalently closed circular dsDNA of 7438 bp (KX008577 = NC_038987) having 3 ORFs, including a large ORF (ORF3) encoding a putative polyprotein with a conserved movement protein, coat protein, aspartic protease, reverse transcriptase, and ribonuclease H domains analogous to those of other badnaviruses (Bomer et al. 2016).

Dioscorea bacilliform SN virus Taxonomic position Genus: Badnavirus

(DBSNV)

Family: Caulimoviridae

Geographical distribution DBSNV infection in plants of Dioscorea sansibarensis was reported from West Africa (Seal and Muller 2007; Eni et al. 2008b). Symptoms and host(s) The virus-infected yam plants show mild leaf deformation and chlorosis symptoms. Transmission The virus is not mechanically sap-transmissible but transmissible through grafting. Virion properties and genome The virions are bacilliform shaped of about 130 nm in length and 30 nm wide. The genome is circular double-stranded DNA of 7261 bp which contains three open reading frames (DQ822073 = NC_009010) (Seal and Muller 2007; Olszewski and Lockhart 2011; Bhat et al. 2016).

Dioscorea bacilliform TR virus Taxonomic position Genus: Badnavirus

(DBTRV)

Family: Caulimoviridae

Dioscorea spp. (Yam)

905

Geographical distribution DBTRV infection in plants of Dioscorea trifida was reported from Guadeloupe (Umber et al. 2017). Transmission The virus-infected planting material is the main source of virus spread. The virus is not mechanically sap-transmissible. Virion properties and genome Virions are bacilliform in shape with a diameter of 30 nm and modal particle length is 130 nm (Olszewski and Lockhart 2011). The genome contains a single molecule of non-covalently closed circular dsDNA of 7333 bp (KX430257 = NC_038995) with 3 ORFs, including a large ORF (ORF3) encoding a putative polyprotein with a conserved movement protein, coat protein, aspartic protease, reverse transcriptase and ribonuclease H domains analogous to those of other badnaviruses (Umber et al. 2017).

Dioscorea latent virus

(DLV)

Taxonomic position DLV is a tentative member of the genus Potexvirus and family Alphaflexiviridae Geographical distribution DLV infection in plants of Dioscorea spp. was first reported from Puerto Rico by Lawson et al. (1973). The virus spreads in Central America, Fiji, and Vanuatu in the South Pacific (Lebas et al. 2002). Symptoms and host(s) The virus-infected Dioscorea composita and, D. floribunda plants do not exhibit any symptoms (Lawson et al. 1973). Transmission The virus is transmissible by mechanical sap-inoculation (Phillips et al. 1986). The virus is not transmissible by contact between plants and by seed and pollen. No vector is known for this virus. Virion Properties and Genome The virions are filaments particles most of which measured c. 350–900 nm in purified preparations, with two modal lengths of 445 and 875 nm. The genome is single-stranded RNA, approximately 7.5 kb in size and a coat protein with a molecular weight of approximately 25 kDa (Phillips et al. 1986).

Dioscorea mosaic associated virus Taxonomic position Genus: Unassigned

(DMaV)

Family: Secoviridae

Geographical distribution DMaV infection in plants of Dioscorea rotundata was reported from Brazil (Hayashi et al. 2017).

D

906

Dioscorea spp. (Yam)

Symptoms and host(s) The virus-infected yam plants exhibit mosaic symptoms. Transmission The virus is transmitted by nematode vectors in non-persistent manner, and also by mechanical sapinoculation, and by grafting. Virion properties and genome Virions are isometric and 25–30 nm in diameter. The virus has a bipartite genome: RNA1 contains 5979 nt (KU215538 = NC_031766) and RNA2 3810 nt (KU215539 = NC_031763) in length, excluding the poly(A) tails. One large open reading frame (ORF) in each genomic segment (RNA1ORF1 and RNA2-ORF2) was predicted, most likely proteolytically cleaved to yield the mature proteins (Hayashi et al. 2017; Thompson et al. 2017).

Japanese yam mosaic virus Taxonomic position Genus: Potyvirus

(JYMV)

Family: Potyviridae

Geographical distribution JYMV was first reported by Okuyama and Saka (1978) from infected Japanese yam (Dioscorea japonica). The virus occurs in certain parts of Asia (Fuji and Nakamae 1999a, b; Kameya-Iwaki et al. 1999; Fuji et al. 2001; Fukuta et al. 2003; Kajihara et al. 2009; Lan et al. 2015; Mochizuki et al. 2017). Symptoms and host(s) The virus-infected yam plants show mosaic and green banding of leaves and result in yield losses. Transmission The virus is transmitted in a non-persistent manner by aphids (Aphis gossypii and Myzus persicae). The virus is mechanically sap-transmissible and is also transmissible through the use of infected planting material (Okuyama and Saka 1978). Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9760 nt (AB027007 = NC_000947) (Fuj and Nakamae 1999a, 2000; Revers and Garcia 2015; Lan et al. 2015; Wylie et al. 2017), which contains a single open reading frame encoding a polyprotein of 3130 amino acids. Previously, this potyvirus was classified as YMV. The complete genome sequence of JYMV showed that it is distinct from YMV, and therefore, JYMV is considered as a separate species under the genus Potyvirus.

Yam chlorotic mosaic virus Taxonomic position Genus: Macluravirus

(YCMV)

Family: Potyviridae

Dioscorea spp. (Yam)

907

Geographical distribution YCMV infection in plants of Dioscorea parviflora and D. zingiberensis was reported from Yunnan Province, China (Wang et al. 2009; Zhang et al. 2016). Symptoms and host(s) The virus-infected plants of D. zingiberensis exhibit combination of vein-clearing, veinal necrosis, and chlorotic symptoms on the expanded leaves, whereas D. parviflora show systemic chlorosis, mottling, and mosaic symptoms that also affect the newly emerging leaves. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically saptransmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, 650–675 nm long, and 13–16 nm in diameter. The genome consists of a single molecule of linear positive-sense ssRNA of 8208 nt (KT724961) (excluding the poly(A) tail) encoding a polyprotein of 2622 amino acids (Lopez-Moya et al. 2009; Foster 2011; Zhang et al. 2016; Wylie et al. 2017).

Yam latent virus Taxonomic position Genus: Carlavirus

(YLV)

Family: Betaflexiviridae

Geographical distribution YLV infection in plants of Dioscorea opposita was reported from China (Zou et al. unpublished; KJ789130). Transmission The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. Virion properties and genome Virions are flexuous filaments about 640 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8531 nt (KJ789130 = NC_026248) and comprises of six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP and a putative nucleic acid-binding regulatory protein. Coat protein subunit is of one type, and 31–36 kDa in size (Zou et al. unpublished; KJ789130).

Yam mild mosaic virus Taxonomic position Genus: Potyvirus

(YMMV)

Family: Potyviridae

D

908

Dioscorea spp. (Yam)

Geographical distribution YMMV infection in plants of Dioscorea spp. was reported from Cote d’Ivoire, China, Guangxi, Indonesia, Solomon Islands, Columbia, Mauritius, Japan, Sri Lanka, Fiji Islands, Benin, Nigeria, Ghana, Togo, Barbados, Martinique, Puerto Rico, Papua New Guinea, Brazil, India, and Columbia (Hughes 1986; Reckhaus 1980; Hughes et al. 1998; Fuji et al. 1999, 2001; Bousalem and Dallot 2000; Dallot et al. 2001; Danmura et al. 2004; Eni et al. 2008c, 2010; Kajihara et al. 2009; Zou et al. 2011; Jeeva et al. 2014; Yeyeh et al. 2014). Symptoms and host(s) This virus often causes mild symptoms on the leaves of the yam and hence the species name Yam mild moaic virus. Variation in symptom seen in certain Dioscorea species was noticed, viz., Dioscorea alata, mild chlorotic mottling; D. dumetorum and D. cayenensis, mosaic; and D. rotundata, chlorotic mottling and mosaic. Transmission The aphid vector Aphis craccivora transmits this disease in a non-persistent manner. The virus is not seed-borne. However, the pathogen is transmitted vegetatively through tubers derived from infected plants. The virus is mechanically sap-transmissible to a number of hosts. Experimental indicator plants of Vigna unguiculata showed chlorosis, bleaching, and green vein-banding. Systemically infected indicator plants of Nicotiana clevelandii, Chenopodium capitatum, and C. quinoa showed no symptoms. Virion properties and genome The virions are non-enveloped, flexuous filaments, 750 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9538 nt (JX470965 = NC_019412) (Odu 1997; Fuji et al. 1999; Filho et al. 2013; Revers and Garcia 2015; Wylie et al. 2017).

Yam mosaic virus

(YMV)

Synonyms Dioscorea alata virus; Dioscorea greenbanding virus; Dioscorea trifida virus; Dioscorea vein banding virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution YMV infection in plants of Dioscorea spp. was first isolated and characterized in the Cote d’Ivoire by Thouvenel and Fauquet (1979). The virus spreads in Africa, Ghana, Togo, the West Indies, Guadeloupe, and Guyana (Terry 1976; Okuyama and Saka 1978; Porth and Nienhaus 1983; Hughes 1986; Porth et al. 1987; Aleman et al. 1996; Goudou-Urbino et al. 1996; Odu et al. 1999; Njukeng et al. 2003, 2014; Bousalem and Loubet 2008; Eni et al. 2008c, 2010; Odedara et al. 2011; Asala et al. 2012; Yeyeh et al. 2014). Symptoms and host(s) The virus-infected Dioscorea spp. plants exhibit symptoms of green mosaic and vein-banding, spotting and downward curling of the leaves, leaf chlorosis, distortion, or shoestringing, and finally lead to

Dioscorea spp. (Yam)

909

stunting of the plants with poor tuber yields. Several strains of YMV were described (Reckhaus and Nienhaus 1981; Porth et al. 1987). Species like Dioscorea alata, D. cayenensis, D. dumetorum, D. praehensilis, D. Preussii, and D. rotundata are susceptible to this disease and exhibit mosaic and vein-banding. Symptoms vary depending on the host species and environmental conditions. Transmission The virus is transmitted by the aphid vectors, Aphis craccivora, A. gossypii, Rhopalosiphum maidis, and Toxoptera citricidus, in a non-persistent manner. The virus is transmissible by mechanical sapinoculation, but the host range is strictly limited to the Dioscoreaceae and Solanaceae. The virus is not transmissible by contact between plants; it is not transmitted by seed (it is by vegetative propagation). The primary means of spread of this disease is use of planting material from an infected source. Virion properties and genome The virions are flexuous filaments, non-enveloped, and with a clear modal length of 785 nm and 15 nm wide. The genome consists of a single-stranded RNA of 9608 nt (U42596 = NC_004752) and contains one open reading frame encoding a polyprotein of 3103 amino acids (aa) (Aleman et al. 1996). The 50 leader sequence of YMV RNA preceding the ORF is 134 nt, while the 30 untranslated region (UTR) is 165 nt excluding the poly(A) tail (Ishikawa et al. 1985; Wylie et al. 2017).

Yam spherical virus Taxonomic position Genus: Aureusvirus

(YSV)

Family: Tombusviridae

Geographical distribution YSV was reported in plants of Dioscorea rotundata from Nigeria (Menzel et al. 2014). Transmission The virus is soil-borne and a soil-inhabiting fungus (Olpidium spp.) is the vector. The virus is mechanically sap-transmissible to Nicotiana benthamiana, resulting in systemic veinal necrosis. The following Nicotiana species are infected systemically: N. hesperis (veinal necrosis), N. rustica ‘NRT 63’ (necrotic spots), N. glutinosa, and N. clevelandii. Inoculation to N. occidentalis “37B” and Cucumis sativus “Vorgebirgstraube” resulted in necrosis on inoculated leaves, whereas Chenopodium quinoa developed necrotic local lesions; no systemic infection of YSV was observed in these three plant species. Virion properties and genome The virions are isometric, non-enveloped, 30 nm in diameter, and have T = 3 icosahedral symmetry. The genome is a monopartite, linear, single-stranded, positive-sense RNA of 4464 nt (KF482072 = NC_022895) (Martelli et al. 1998; Martelli and Rubino 2011; Menzel et al. 2014).

Yam virus X Taxonomic position Genus: Potexvirus

(YVX)

Family: Alphaflexiviridae

D

910

Dioscorea spp. (Yam)

Geographical distribution YVX infection in plants of Dioscorea spp. was reported from Guadeloupe (Mambole et al. 2014). Symptoms and host(s) The virus-infected yam plants exhibit mild mosaic symptoms. Transmission The virus is mechanically sap-transmissible. No vector is known for this virus. Virion properties and genome The virions are flexuous filaments, 470–580 nm in length, and 13 nm in diameter. The genome is a linear single molecule of positive-sense single-stranded RNA of 6158 nt (KJ711908) and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type, 18–27 kDa in size (Adams et al. 2004; Mambole et al. 2014).

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Aleman ME, Marcos JF, Brugidou C, Beachy RN, Fauquet C (1996) The complete nucleotide sequence of Yam mosaic virus (Ivory Coast isolate) genomic RNA. Arch Virol 141:1259–1278 Asala S, Alegbejo MD, Kashina B, Banwo OO, Asiedu R, Lava-Kumar P (2012) Distribution and incidence of viruses infecting yam (Dioscorea spp.) in Nigeria. GJBB 1:163–167 Bhat AI, Hohn T, Selvarajan R (2016) Badnaviruses: the current global scenario. Viruses 8(6):177 Bomer M, Turaki A, Silva G, Kumar P, Seal SA (2016) Sequence-independent strategy for amplification and characterisation of episomal Badnavirus sequences reveals three previously uncharacterised yam badnaviruses. Viruses 8:188 Bousalem M, Dallot S (2000) First report and molecular characterization of Yam mild mosaic virus in Dioscorea alata on the island of Martinique. Plant Dis 84:200 Bousalem M, Loubet S (2008) Molecular evidence for a new potyvirus species in yam (Dioscorea spp.) on the island of Guadeloupe. Plant Pathol 57:389 Briddon RW, Phillips S, Brunt A, Hull R (1999) Analysis of the sequence of Dioscorea alata bacilliform virus: comparison to other members of the badnavirus. Virus Genes 18:277–283 Dallot S, Guzman M, Bousalem M (2001) Occurrence of potyviruses on yam (Dioscorea spp.) in Colombia and first molecular characterization of Yam mild mosaic virus. Plant Dis 85:803 Danmura K, Ito S, Tanaka S, Kameya-Iwaki M, Kajihara H, Fuji S (2004) Detection of yam mild mosaic virus from Dioscorea plants by immunocapture reverse transcription-polymerase chain reaction. Jpn J Phytopathol 70:264 Dey KK, Sugikawa J, Kerr C, Melzer MJ (2019) Air potato (Dioscorea bulbifera) plants displaying virus-like symptoms are co-infected with a novel Potyvirus and a novel Ampelovirus. Virus Genes 55(1):117–121 Eni AO, Hughes J’A, Rey MEC (2008a) First report of Cucumber mosaic virus in yam (Dioscorea spp.) in Ghana, Togo and Republic of Benin in West Africa. Plant Dis 92:833 Eni AO, Jd H, Asiedu R, Rey ME (2008b) Sequence diversity among badnavirus isolates infecting yam (Dioscorea spp.) in Ghana, Togo, Benin and Nigeria. Arch Virol 153(12):2263–2272 Eni AO, Hughes JA, Rey MEC (2008c) Survey of the incidence and distribution of five viruses infecting yams in the major yam-producing zones in Benin. Ann Appl Biol 153:223–232 Eni AO, Hughes J’A, Asiedu R, Rey MEC (2010) Survey of the incidence and distribution of viruses infecting yam (Dioscorea spp.) in Ghana and Togo. Ann Appl Biol 156:243–251 Eni AO, Lava Kumar P, Asiedu R, Alabi OJ, Naidu RA, Hughes J’A, Rey MEC (2013) Characterization of Cucumber mosaic virus isolated from yam (Dioscorea spp.) in West Africa. Afr J Biotechnol 12:3472–3480 Fauquet C, Thouvenel JC (1987) Plant viral diseases in the Ivory Coast. ORSTOM Paris Doc Tech 46:243 Filho FD, Nicolini C, Resende RD, Andrade GP, Pio-Ribeiro G, Nagata T (2013) The complete genome sequence of a Brazilian isolate of yam mild mosaic virus. Arch Virol 158(2):515–518 Foster GD (2011) Macluravirus. Potyviridae. The Springer index of viruses. Springer, New York, pp 1421–1424. https:// doi.org/10.1007/978-0-387-95919-1_234

Dioscorea spp. (Yam)

911

Fuji S, Nakamae H (1999a) Complete nucleotide sequence of the genomic RNA of a Japanese yam mosaic virus, a new potyvirus in Japan. Arch Virol 144:231–240 Fuji S, Nakamae H (1999b) Detection of Japanese yam mosaic virus by ELISA and RT-PCR. Ann Phytopathol Soc Jpn 65:207–210 Fuji S, Nakamae H (2000) Complete nucleotide sequence of the genomic RNA of a mild strain of Japanese yam mosaic potyvirus. Arch Virol 145(3):635–640 Fuji S, Mitobe I, Nakamae H, Natsuaki KT (1999) Nucleotide sequence of coat protein gene of Yam mild mosaic virus, isolated in Papua New Guinea. Arch Virol 144:1415–1419 Fuji S, Furuya H, Naito H, Natsuaki KT (2001) Detection of potyviruses in yam plants cultivated in Japan. Jpn J Phytopathol 67:261–263 Fukuta S, Iida T, Mizukami Y, Ishida A, Ueda J, Kanbe M, Ishimoto Y (2003) Detection of Japanese yam mosaic virus by RT-LAMP. Arch Virol 148:1713–1720 Fukumoto F, Tochihara H (1978) Chinese yam necrotic mosaic virus. Ann Phytopath Soc Jpn 44:1–5 Goudou-Urbino C, Givord L, Konate G, Boeglin M, Quiot JB, Dubern J (1996) Differentiation of yam virus isolates by using symptomatology, western-blot and monoclonal antibodies. J Phytopathol 144:235–240 Harrison BD, Roberts IM (1973) Association of virus-like particles with internal brown spot of yam (Dioscorea alata). Trop Agric 50:335–340 Hayashi EAI, Blawid R, de Melo FL, Andrade MS, Pio-Ribeiro G, de Andrade GP, Nagata T (2017) Complete genome sequence of a putative new secovirus infecting yam (Dioscorea) plants. Arch Virol 162(1):317–319 Hughes Jd’A (1986) Viruses of Araceae and Dioscoreaceae: their isolation, characterization and detection. PhD thesis, University of Reading, 160 p Hughes J’A, Dongo L, Atiri GI (1997) Viruses infecting cultivated yams (Dioscorea alata and D. rotunda) in Nigeria. Phytopathol 87:545 Hughes J’A, Dongo LN, Ng SYC (1998) Diagnosis of yam viruses. Tropi Agric Trinidad 75:45–48 Ishikawa R, Nieta H, Namba S, Yamashita S, Doi Y (1985) Purification of a small spherical virus and Yam mosaic virus in Chinese yam. Anna Phytopathol Soc Jpn 51:99 Jeeva ML, Makeshkumar T, Rajitha M, Manasa VG, Sruthy S (2014) Investigation of greater yam (Dioscorea alata L.) viruses in India. In: Virocon 2014- IVS- XXIII national conference on recent trends in virology research in the Omics era, held at TNAU, Coimbatore during 18–20 December 2014, p 68 Kajihara H, Muramoto K, Fuji S, Tanaka S, Ito S (2009) Simultaneous detection of Japanese yam mosaic virus and Yam mild mosaic virus from yam leaves using a tube capture reverse transcription-polymerase chain reaction assay. J Gen Plant Pathol 75:72–75 Kameya-Iwaki M, Yamaguchi K, Hara T, Ito S, Fuji S, Kajihara H, Tanaka S (1999) Detection of Japanese yam mosaic virus from Chinese yam (dioscorea opposita thumb. Cv. Ichoimo) by immunocapture PCR and differentiation of virulent and attenuated isolates by PCR-RFLP analysis. Ann Phytopathol Soc Jpn 65:494–497 Kondo T (2001) The 30 terminal sequence of Chinese yam necrotic mosaic virus genomic RNA: a close relationship with macluravirus. Arch Virol 146:1527–1535 Kondo T, Fujita T (2012) Complete nucleotide sequence and construction of an infectious clone of Chinese yam necrotic mosaic virus suggest that macluraviruses have the smallest genome among members of the family Potyviridae. Arch Virol 157:2299–2307 Kondo T, Kang DK, Fuji S, Chang MU (2003) Sequence diversity in the coat protein and 30 -untranslated region of Chinese yam necrotic mosaic virus RNA. Jo Gen Plant Pathol 69:397–399 Kondo T, Fuji S, Yamashita K, Kang D-K, Chang M-U (2005) Broad bean wilt virus 2 in yams. J Gen Plant Pathol 71:441–443 Kwon SJ, Choi S, Yoon JY, Choi SK, Choi GS (2016) First report of broad bean wilt virus 2 in Dioscorea opposita thumb. In Korea. Plant Dis 100:538 Lan P, Li F, Wang M, Li R (2015) Complete genome sequence of a divergent strain of Japanese yam mosaic virus from China. Arch Virol 160:573–576 Lawson RH, Haron SS, Smith FF, Kahn RP (1973) Electron microscopy and separation of viruses in Dioscorea floribunda. Phytopathology 63:1435 Lebas B, Canning E, Kenyon L, Seal S (2002) Yam viruses of the South Pacific Islands. Pest Management Department, University of Greenwich, Natural Resources Institute, UK Lopez-Moya JJ, Valli A, Garcia JA (2009) Potyviridae. In: Encyclopaedia of life sciences (ELS). Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000755.pub2 Mambole IA, Bonheur L, Dumas LS, Filloux D, Gomez RM, Faure C, Lange D, Anzala F, Pavis C, Marais A, Roumagnac P, Candresse T, Teycheney PY (2014) Molecular characterization of yam virus X, a new potexvirus infecting yams (Dioscorea spp) and evidence for the existence of at least three distinct potexviruses infecting yams. Arch Virol 159:3421–3426 Martelli GP, Rubino L (2011) Aureusvirus. Tombusviridae. The Springer index of viruses. Springer, New York, pp 1162–1164. https://doi.org/10.1007/3-540-31042-8_200

D

912

Dioscorea spp. (Yam)

Martelli GP, Russo M, Rubino L, Sabanadzovic S (1998) Aureusvirus, a novel genus in the family Tombusviridae. Arch Virol 143:1847–1851 Menzel W, Thottappilly G, Winter S (2014) Characterization of an isometric virus isolated from yam (Dioscorea rotundata) in Nigeria suggests that it belongs to a new species in the genus Aureusvirus. Arch Virol 159:603–606 Migliori A, Cadilhac B (1976) Contribution A L’etude de la maladie du virus de I’igname Dioscorea trifida en Guadeloupe. Ann Phytopathol 8:73–78 Mochizuki T, Iwamoto I, Atarashi A, Ohki ST, Kikukawa H (2017) Rapid and low-cost diagnosis of Japanese yam mosaic virus infection in Chinese yam (Dioscorea polystachya) leaves by a print-capture RT-PCR. J Gen Plant Pathol 83:189–196 Njukeng AP, Hughes Jd’A, Atiri GL, Kpo, EJA (2003) Distribution of Yam mosaic virus in Nigeria 8th Intern. Plant Virus Epidemiology Symposium. Aschersleben, 12–17 Njukeng AP, Azeteh IN, Mbong GA (2014) Survey of the incidence and distribution of two viruses infecting yam (Dioscorea spp) in two agro-ecological zones of Cameroon. Int J Curr Microb App Sci 3:1153–1166 Odedara OO, Ayo-John EI, Gbuyiro MM, Falade FO, Agbebi SE (2011) Serological detection of yam viruses in farmers’ fields in Ogun state, Nigeria, arch. Phytopathol Plant Protect 45(7):840–845 Odu BO (1997) Characterization of a yam potyvirus in Dioscorea alata L. (water yam). MSc thesis. University of Ibadan, Nigeria Odu BO, Hughes J’A, Shoyinka SA, Dongo LN (1999) Isolation, characterization and identification of potyvirus from Dioscorea alata (water yam) in Nigeria. Ann Appl Biol 134:65–71 Okuyama S, Saka H (1978) Yam mosaic virus. Sci Rep Fac Agric Ibaraki Univ 26:29–34 Olszewski NE, Lockhart B (2011) Badnavirus. Caulimoviridae. The Springer index of viruses. Springer, New York, pp 263–269. https://doi.org/10.1007/978-0-387-95919-1_40 Phillips S, Piggott JD’A, Brunt AA (1986) Further evidence that Dioscorea latent virus is a Potexvirus. Ann Appl Biol 109:137–145 Phillips S, Briddon RW, Brunt AA, Hull R (1999) The partial characterization of a badnavirus infecting the greater Asiatic or water yam (Dioscorea alata). J Phytopathol 147:265–269 Porth A, Nienhaus F (1983) Dioscorea alata ring mottle virus, a new potyvirus of yam in Togo. Z Pfl Krankh Pfl Schutz 90:352–362 Porth A, Lesemann DE, Vetten HJ (1987) Characterization of potyvirus isolates from West African Yams (Dioscorea spp.). J Phytopathol 90:352–362 Reckhaus P (1980) Untersuchungen zur Atiologie virus Verdachtiger arkenkungen an yam (Dioscorea spp.) in West Africa (Togo). PhD thesis, University of Bonn, 103 p Reckhaus P, Nienhaus F (1981) Etiology of a virus disease of white yam D. rotundata in Togo. J Plant Dis Protect 88:492–509 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Seal S, Muller E (2007) Molecular analysis of a full-length sequence of a new yam badnavirus from Dioscorea sansibarensis. Arch Virol 152:819–825 Sukal A, Kidanemariam D, Dale J, James A, Harding R (2017) Characterization of badnaviruses infecting Dioscorea spp. in the Pacific reveals two putative novel species and the first report of Dioscorea bacilliform RT virus 2. Virus Res 238:29–34 Terry ER (1976) Incidence, symptomatology, and transmission of a yam virus in Nigeria. In: Cock J, McIntyre and Graham M (eds) Proceedings of the Fourth Symposium of the International Society for Tropical Root Crops, CIAT, Cali, August 1976 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Secoviridae. J Gen Virol 98:529–531 Thouvenel JC, Fauquet C (1979) Yam mosaic, a new potyvirus infecting Dioscorea cayenensis in the ivory coast. Ann Appl Biol 93:279–283 Umber M, Gomez RM, Gélabale S, Bonheur L, Pavis C, Teycheney PY (2017) The genome sequence of Dioscorea bacilliform TR virus, a member of the genus Badnavirus infecting Diocorea spp., sheds light on the possible function of endogenous Dioscorea bacilliform viruses. Arch Virol 162(2):517–521 Wang JG, Zou XJ, Zheng HY, Adams MJ, Chen HR, Chen JP (2009) Molecular characterization of a new macluravirus from yam in Yunnan, China. Arch Virol 154:1379–1380 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354 Yeyeh TMN, Atta Diallo H, Akinbade SA, Seka K, Lava Kumar P (2014) Distribution, incidence and severity of viral diseases of yam (Dioscorea spp.) in cote d’Ivoire. Afr J Biotechnol 13:465–470 Zhang P, Peng J, Guo H, Chen J, Chen S, Wang J (2016) Complete genome sequence of yam chlorotic necrotic mosaic virus from Dioscorea parviflora. Arch Virol 161(6):1715–1717 Zou C, Meng J, Li Z, Wei M, Song J, Chen B, Wei B (2011) First report of Yam mild mosaic virus in yam in Guangxi province, China. Plant Dis 95:1320

Diospyros kaki (Japanese persimmon)

913

Diospyros kaki (Japanese persimmon) Family: Ebenaceae

Edible fruit

Apple fruit crinkle viroid

(AFCVd)

Taxonomic position AFCVd is a tentative member of the genus Apscaviroid and family Pospiviroidae An isolate of AFCVd was detected in plants of Diospyros kaki in Japan (Nakaune and Nakano 2008). The virioid-infected Japanese persimmon plants exhibit fruit apex disorder symptoms. The viroid is easily transmissible by budding, grafting, and chip budding and was transmissible mechanically. For more details of AFCVd, refer to Malus domestica.

Citrus viroid VI

(CVd VI)

Synonyms Citrus viroid OS (CVd-OS) Taxonomic position Genus: Apscaviroid

Family: Pospiviroidae

An isolate of CVd-OS was detected in plants of Diospyros kaki in Japan (Nakaune and Nakano 2008). The viroid-infected Japanese persimmon plants exhibit symptoms of fruit apex disorder that causes necrosis of sieve tissue under the fruit skin. The viroid is transmissible by grafting, cuttings, and mechanical sap-inoculation. For more details of CVd VI, refer to Citrus spp.

Persimmon cryptic virus

(PeCV)

Taxonomic position The virus is a tentative member of the genus Deltapartitivirus and family Partitiviridae Geographical distribution PeCV infection in plants of Diospyros kaki was reported from Italy, Turkey, and Spain (Morelli et al. 2012; 2015; Cho et al. 2016; Morelli and Arli-Sokmen 2016; Ruiz-Garcia et al. 2017). Symptoms and host(s) The virus-infected Japanese persimmon plants exhibit interveinal chlorosis and subsequently lead to extensive necrosis of the veinlets on both sides of the leaf blades. Virion properties and genome The genome consists of two molecules of linear double-stranded RNA. RNA1 contains 1577 nt (HE805113) and RNA2 of 1491 nt (HE805114) (Morelli et al. 2015; Vainio et al. 2018).

D

914

Diospyros kaki (Japanese persimmon)

Persimmon latent viroid

(PLVd)

Synonyms Persimmon viroid (PVd) Taxonomic position PLVd is a tentative member of the genus Apscaviroid and family Pospiviroidae Geographical distribution PLVd was reported in plants of Diospyros kaki from Japan (Nakaune and Nakano 2008). Symptoms and host(s) The Japanese persimmon trees were suffering from graft-transmissible fruit apex disorder that causes necrosis of sieve tissue under the fruit skin. Transmission The viroid is graft-transmissible from persimmon to persimmon. Etiology and genome properties The viroid consists of 396 nt (NC_010308). The predicted secondary structure is rodlike with extensive base pairing which contains the terminal conserved region and the central conserved region characteristic of the genus Apscaviroid.

Persimmon virus A

(PeVA)

Taxonomic position PeVA is a tentative member of the genus Cytorhabdovirus and family Rhabdoviridae Geographical distribution PeVA infection in plants of Diospyros kaki was reported from Italy, Japan, and Korea (Ito et al. 2013; Morelli et al. 2014; Cho et al. 2016). Symptoms and host(s) The virus-infected Japanese persimmon plants exhibit symptoms of extensive necrosis of the veinlets on both sides of the leaf blades. Virion properties and genome The genome is negative-sense single-stranded RNA of 13,467 nt (AB735628) (Ito et al. 2013; Walker et al. 2018).

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

Diospyros virginiana (American persimmon)

915

TRV infection in plants of Diospyros kaki was reported from Egypt (Zein and Salwa 2004). The virusinfected Japanese persimmon plants exhibit symptoms of vein-banding, upward rolling of leaves, and stunting of growth. The virus is transmitted by nematode vectors and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

D

References Cho IS, Park MJ, Kwon SJ, Choi GS, Hammond J, Lim HS (2016) First report of persimmon cryptic virus and persimmon virus A in Korea. J Plant Pathol 98:694 Ito T, Suzaki K, Nakano M (2013) Genetic characterization of novel putative rhabdovirus and dsRNA virus from Japanese persimmon. J Gen Virol 94:1917–1921 Morelli M, Arli-Sokmen M (2016) First report of Persimmon cryptic virus in Turkey. J Plant Pathol 98(1):183 Morelli M, De Stradis A, La Notte P, Merkuri J, Boscia D, Minafra A (2012) Detection and molecular characterization of a novel Cryptovirus from persimmon (Diospyros kaki). Petria 22(3):181 Morelli M, Chiumenti M, La Notte P, Minafra A, Martelli GP (2014) First report of persimmon virus A in Italy. J Plant Pathol 96:610 Morelli M, Chiumenti M, De Stradis A, La Notte P, Minafra A (2015) Discovery and molecular characterization of a new Cryptovirus dsRNA genome from Japanese persimmon through conventional cloning and high-throughput sequencing. Virus Genes 50(1):160–164 Nakaune R, Nakano M (2008) Identification of a new Apscaviroid from Japanese persimmon. Arch Virol 153:969–972 Ruiz-Garcia AB, Chamberland N, Martinez C, Massari S, Olmos A (2017) First report of Persimmon cryptic virus in Spain. J Plant Pathol 99(1):287 Vainio EJ, Chiba S, Ghabrial SA, Maiss E, Roossinck M, Sabanadzovic S, Suzuki N, Xie J, Nibert M, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Partitiviridae. J Gen Virol 99:17–18 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Rhabdoviridae. J Gen Virol 99:447–448 Zein, Salwa N (2004) Characterization of tobacco rattle Tobravirus from Kaki (Diospyros kaki). Egyption J Virol 1:187–193

Diospyros virginiana (American persimmon) Family: Ebenaceae

Persimmon virus B Taxonomic position Genus: Unassigned

Edible fruit

(PeVB)

Family: Closteroviridae

Geographical distribution PeVB infection in plants of Diospyros virginiana was reported from Japan (Ito et al. 2015). Symptoms and host(s) The virus-infected American persimmon plants exhibit leaf petiole and midvein declining symptoms.

916

Diplotaxis erucoides (White rocket)

Virion properties and genome The virions are flexuous filaments, 12 nm wide and length ranging from 650 nm to over 2000 nm. The genome is positive-sense single-stranded RNA of 18,569 nt (AB923924) with 11 open reading frames (Ito et al. 2015). The 50 and 30 UTRs are 183 nt and 282 nt, respectively.

Persimmon viroid 2

(PVd 2)

Taxonomic position PVD 2 is a tentative member of the genus Apscaviroid and family Pospiviroidae Geographical distribution PVd 2 infection in plants of Diospyros virginiana was reported from Japan (Ito et al. 2013). Symptoms and host(s) The viroid-infected American persimmon plants exhibit poor rootstock development and declining symptoms (Ito et al. 2013). Transmission The viroid is transmissible by grafting. Etiology and genome properties The viroid consists of 358 nt (NC_021720) (Ito et al. 2013).

References Ito T, Suzaki K, Nakano M, Sato A (2013) Characterization of a new apscaviroid from American persimmon. Arch Virol 158:2629–2631 Ito T, Sato A, Suzaki K (2015) An assemblage of divergent variants of a novel putative closterovirus from American persimmon. Virus Genes 51:105–111

Diplotaxis erucoides (White rocket) Family: Brassicaceae

Weed host

Pelargonium zonate spot virus Taxonomic position Genus: Anulavirus

(PZSV)

Family: Bromoviridae

PZSV infection in plants of Diplotaxis erucoides was reported from Italy (Gallitelli et al. 2005). Thrips are involved in transmitting this virus by carrying the virus-infected pollen grains on their bodies and introducing the virus into the susceptible hosts while feeding. The virus is transmissible by mechanical sap-inoculation to a wide range of test plants. The virus is also seed-transmitted in D. erucoides with an efficiency of 5% (Vovlas et al. 1989). For more details of PZSV, refer to Pelargonium spp.

Dipsacus fullonum (Fuller’s Teasel)

Pepino mosaic virus

917

(PepMV)

Taxonomic position Genus: Potexvirus

Family: Alphaflexiviridae

PepMV infection in plants of Diplotaxis erucoides was reported from Spain (Cordoba et al. 2004). The virus is mechanically sap-transmissible. For more details of PepMV, refer to Solanum muricatum.

D References Cordoba MC, Martinez-Priego LI, Jorda C (2004) New natural hosts of Pepino mosaic virus in Spain. Plant Dis 88:906 Gallitelli D, Finetti-Sialer M, Martelli GP (2005) Anulavirus, a proposed new genus of plant viruses in the family Bromoviridae. Arch Virol 150:407–411 Vovlas C, Gallitelli D, Conti M (1989) Preliminary evidence for an unusual mode of transmission in the ecology of Pelargonium zonate spot virus (PZSV). Proceedings of 4th plant virus epidemiology workshop, Montpellier, France, pp 302–305

Diplotaxis tenuifolia (Perennial wall-rocket) Family: Brassicaceae

Medicinal plant

Parietaria mottle virus Taxonomic position Genus: Ilarvirus

(PMoV)

Family: Bromoviridae

PMoV infection in plants of Diplotaxis tenuifolia was reported from Italy (Parrella et al. 2017). The virus-infected perennial wall-rocket plants exhibit yellowing symptoms. The virus is transmitted by thrips vectors, and the virus is also mechanically sap-transmissible. For more details of PMoV, refer to Parietaria officinalis.

References Parrella G, Greco B, Troiano E (2017) First report of Parietaria mottle virus associated with yellowing disease in Diplotaxis tenuifolia in Italy. Plant Dis 101:850

Dipsacus fullonum (Fuller’s Teasel) Synonyms Dipsacus sylvestris Family: Caprifoliaceae

Ornamental

918

Dittrichia viscosa (False yellowhead)

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Dipsacus sylvestris was reported from northern Italy (EU432179) (Davino et al. 2012). The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Sunflower chlorotic mottle virus Taxonomic position Genus: Potyvirus

(SCMoV)

Family: Potyviridae

SCMoV infection in plants of Dipsacus fullonum was reported from Argentina (Stoner 1951; Giolitti et al. 2009). The virus-infected fuller’s teasel plants exhibit foliar mosaic symptoms. The virus is transmitted by aphids in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of SCMoV, refer to Helianthus annuus.

References Davino S, Panno S, Rangel EA, Davino M, Bellardi MG, Rubio L (2012) Population genetics of cucumber mosaic virus infecting medicinal, aromatic and ornamental plants from northern Italy. Arch Virol 157:739–745 Giolitti F, Bejerman N, Lenardon S (2009) Dipsacus fullonum: an alternative host of sunflower chlorotic mottle virus in Argentina. J Phytopathol 157:325–328 Stoner WN (1951) An aphid-transmitted virus disease of Fuller’s teasel and pincushion flower. Phytopathology 41:191–194

Dittrichia viscosa (False yellowhead) Family: Asteraceae

Weed host

Tomato infectious chlorosis virus Taxonomic position Genus: Crinivirus

(TICV)

Family: Closteroviridae

TICV infection in plants of Dittrichia viscosa was reported from Greece (Orfanidou et al. 2016). The virus-infected false yellowhead plants exhibit yellowing symptoms. The virus is transmitted by the whitefly vector, Trialeurodes vaporariorum, in a semi-persistent manner. The virus is not transmissible by mechanical sap-inoculation. For more details of TICV, refer to Solanum lycopersicum.

Diuris spp. (Donkey orchid)

919

Reference Orfanidou CG, Maliogka VI, Katis NI (2016) False yellowhead (Dittrichia viscosa), a banker plant as source of tomato infectious chlorosis virus in Greece. Plant Dis 100:869

Diuris spp. (Donkey orchid) Family: Orchidaceae

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

D

Ornamental

(BYMV)

Family: Potyviridae

BYMV was reported from plants of Diuris magnifica and D. corymbosa orchids from Australia (Wylie et al. 2013a). The virus-infected donkey orchid plants exhibit symptoms of flower spike distortion, necrotic patches on leaves, and chlorotic mottling. The virus is transmitted by aphid vectors in a nonpersistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Blue squill virus A Taxonomic position Genus: Potyvirus

(BSVA)

Family: Potyviridae

BSVA infection in plants of Diuris corymbosa was reported from Australia (Wylie et al. 2013a). The virus-infected donkey orchid plants exhibit mild mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BSVA, refer to Chamaescilla corymbosa.

Diuris pendunculata cryptic virus

(DPCV)

Taxonomic position DPCV is a tentative member of the genus Alphapartitivirus and family Partitiviridae Geographic distribution DPCV infection in plants of Diuris pendunculata was reported from Western Australia (Wylie et al. 2013a). Symptoms and host(s) The virus-infected donkey orchid plants do not exhibit visible symptoms.

920

Diuris spp. (Donkey orchid)

Virion properties and genome Virions are isometric, 25–50 nm in diameter. The genome consists of double-stranded RNA of two parts. RNA1 contains 2010 bp (JX156424) and RNA2 of 1806 bp (JX891460) (Vainio et al. 2018).

Diuris virus A

(DiV-A)

Taxonomic position Genus: Divavirus

Family: Betaflexiviridae

Geographical distribution DiV-A infection in plants of Diuris laxiflora was reported from Australia (Wylie et al. 2013a). Symptoms and host(s) The virus-infected donkey orchid plants do not show visible symptoms. Virion properties and genome The genome is a single-stranded RNA of 6956 nt (NC_019029) (Wylie et al. 2013a).

Diuris virus B Taxonomic position Genus: Divavirus

(DiV-B)

Family: Betaflexiviridae

Geographical distribution DiV-B infection in plants of Diuris pendunculata was reported from Western Australia (Wylie et al. 2013a). Symptoms and host(s) The virus-infected donkey orchid plants do not exhibit visible symptoms. Virion properties and genome The genome is a single-stranded RNA of 7001 nt (NC_019030).

Diuris virus Y

(DiV-Y)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution DiV-Y was reported from plants of Diuris orchids from Australia (Gibbs et al. 2000). Symptoms and host(s) The virus-infected donkey orchid plants exhibit symptoms of mottling on leaves.

Diuris spp. (Donkey orchid)

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Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome is a single molecule of linear, positive-sense, single-stranded RNA. A partial genome sequence of 1719 nt is available (AF203527) (Gibbs et al. 2000; Wylie et al. 2017).

Donkey orchid symptomless virus Taxonomic position Genus: Platypuvirus

(DOSV)

Family: Alphaflexiviridae

Geographical distribution DOSV infection in plants of Diuris longifolia was reported from Australia (Wylie et al. 2013b). Symptoms and host(s) The virus-infected donkey orchid plants do not exhibit any symptoms. The virus was also identified in Caladenia latifolia (pink fairy orchid) and from Drakaea elastica. Transmission The virus is mechanically sap-transmissible. Virion properties and genome The virions are flexuous. The positive-sense single-stranded RNA genome of 7838 nt (KC923234 = NC_022894) has an unusual organization, unlike that of other classified viruses, and has a predicted 30 pseudoknot and no poly(A) tract (Wylie et al. 2013b).

Donkey orchid virus A Taxonomic position Genus: Potyvirus

(DOVA)

Family: Potyviridae

Geographic distribution DOVA infection in plants of Diuris laxiflora was reported from Western Australia (Wylie et al. 2013a). Symptoms and host(s) The virus-infected donkey orchid plants do not show visible symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically sap-transmissible.

D

922

Dolichos lablab (Hyacinth bean)

Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9868 nt (JX156422 = NC_021197) (Wylie et al. 2017).

Ornithogalum mosaic virus Taxonomic position Genus: Potyvirus

(OrMV)

Family: Potyviridae

OrMV infection in plants of Diuris spp. was reported from Western Australia (Wylie et al. 2013a). The virus-infected donkey orchid plants exhibit symptoms of chlorotic leaf mottle and distortion of the flower spike. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of OrMV, refer to Ornithogalum spp.

Turnip yellows virus Taxonomic position Genus: Polerovirus

(TuYV)

Family: Luteoviridae

TuYV infection in plants of Diuris pendunculata was reported from Western Australia (Wylie et al. 2013a). The virus is transmitted by a number of aphid species in a circulative and non-propagative manner. The virus is not mechanically sap-transmissible. For more details of TuYV, refer to Brassica rapa.

References Gibbs A, Mackenzie A, Blanchfield A, Cross P, Wilson C, Kitajima EW, Nightingale M, Clements M (2000) Viruses of orchids in Australia: their identification, biology and control. Aust Orchid Rev 65:10–21 Vainio EJ, Chiba S, Ghabrial SA, Maiss E, Roossinck M, Sabanadzovic S, Suzuki N, Xie J, Nibert M, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Partitiviridae. J Gen Virol 99:17–18 Wylie SJ, Li H, Dixon KW, Richards H, Jones MGK (2013a) Exotic and indigenous viruses infect wild populations and captive collections of temperate terrestrial orchids (Diuris species) in Australia. Virus Res 171:22–32 Wylie SJ, Li H, Jones MGK (2013b) Donkey orchid symptomless virus: a viral ‘platypus’ from Australian terrestrial orchids. PLoS One 8(11):e79587. https://doi.org/10.1371/journal.pone.0079587 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354

Dolichos lablab (Hyacinth bean) Synonyms Lablab purpureus Family: Fabaceae

Grain legume

Dolichos lablab (Hyacinth bean)

923

Bean common mosaic necrosis virus Taxonomic position Genus: Potyvirus

(BCMNV)

Family: Potyviridae

BCMNV was reported in plants of Lablab purpureus from India, Uganda, and Nepal (Sengooba et al. 1997; Pudashini et al. 2013; Pavithra et al. 2017). The virus-infected hyacinth bean plants exhibit symptoms of mottle and leaf deformation, severe mosaic, necrosis, malformation of leaves and pods, downward curling of leaves, and reduction in leaf size. The virus is transmitted by an aphid vector, Aphis fabae, in a non-persistent manner, and is also transmissible by mechanical inoculation. For more details of BCMNV, refer to Phaseolus vulgaris.

Bean common mosaic virus Taxonomic position Genus: Potyvirus

(BCMV)

Family: Potyviridae

BCMV infection in plants of Lablab purpureus was reported from India and Nigeria (Odedara et al. 2008; Udayashankar et al. 2011). The virus-infected hyacinth bean plants exhibit symptoms of stunting, mosaic, vein-banding, vein-clearing, mottling, and blisters. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BCMV, refer to Phaseolus vulgaris.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Lablab purpureus was reported from India (Rishi and Dhawan 1987). The virus-infected hyacinth bean plants exhibit symptoms of discolored areas on leaves with distinct bright yellow patches. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Clover yellow vein virus Taxonomic position Genus: Potyvirus

(ClYVV)

Family: Potyviridae

ClYVV infection in plants of Dolichos lablab was reported from China (Li 1991). The virus-infected hyacinth bean plants exhibit systemic mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ClYVV, refer to Trifolium spp.

D

924

Dolichos lablab (Hyacinth bean)

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Dolichos lablab was reported from China, India, and Nigeria (Zhou and Chen 1990; Kiranmai et al. 1998; Odedara et al. 2008). The virus-infected hyacinth bean plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Dolichos yellow mosaic virus Taxonomic position Genus: Begomovirus

(DoYMV)

Family: Geminiviridae

Geographical distribution DoYMV infection in plants of Dolichos lablab was reported from India and Pakistan (Capoor and Varma 1950; Subramanian 1975; Raj et al. 1988; Maruthi et al. 2006a, b; Singh et al. 2012). Symptoms and host(s) The virus-infected hyacinth bean plants exhibit yellow mosaic symptoms. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci. Insects required a minimum of 6 h to acquire the virus and 1 h to transmit the virus. By insect transmission the virus could only be transferred to Lablab purpureus (Maruthi et al. 2006b). The virus is also transmissible by grafting. Virion properties and genome Non-enveloped virions with a geminate (twinned quasi-isometric) morphology of ~20  30 have been shown to be associated with Dolichos lablab plants with yellow mosaic symptoms (Raj et al. 1988). The structure of the virions of DoYMV has not been investigated. Monoclonal antibodies raised against African cassava mosaic virus and Indian cassava mosaic virus cross react with DoYMV (Maruthi et al. 2006a, b). The genome of DoYMV is bipartite consisting of two circular, single-stranded DNA components. DNA-A contains 2761 nt (KJ481204; AY271891) and DNA-B of 2733 nt (KJ481205) (Raj et al. 1988; Briddon 2001; Qazi et al. 2007; Brown et al. 2015; Akram et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the Old World, the DNA-A component of DoYMV encodes six genes, two in the virion-sense and four in the complementary-sense, and two of the DNA-B component, one in each orientation. The expression and function of these genes has not been investigated for DoYMV.

Peanut stunt virus Taxonomic position Genus: Cucumovirus

(PSV)

Family: Bromoviridae

Dolichos lablab (Hyacinth bean)

925

PSV infection in plants of Dolichos lablab was reported from Sudan (Ahmed and Mills 1985). The virus-infected hyacinth bean plants exhibit bright yellow mottle symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of PSV, refer to Arachis hypogaea.

Soybean mosaic virus Taxonomic position Genus: Potyvirus

(SMV)

D Family: Potyviridae

SMV infection in plants of Dolichos lablab was reported from China (Zhou and Chen 1990). The virusinfected hyacinth bean plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of SMV, refer to Glycine max.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Lablab purpureus was reported from Andhra Pradesh (India) (Bhaskara Reddy et al. 2014). The virus-infected hyacinth bean plants exhibit symptoms of leaf veinal necrosis and necrotic spotting and necrotic streaks on stems and petioles. The virus is transmitted by the thrips vectors, the virus present in/on pollen, and enters the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible but is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

References Ahmed AH, Mills PR (1985) Identification of peanut stunt virus in the Sudan. Plant Dis 69:173–174 Akram M, Naimuddin AAK, Gupta S, Singh NP (2015) Characterisation of full genome of Dolichos yellow mosaic virus based on sequence comparison, genetic recombination and phylogenetic relationship. Ann Appl Biol 167:354–363 Bhaskara Reddy BV, Prasanthi L, Sivaprasad Y, Sujitha A, Giridhar Krishna T (2014) First report of the natural occurrence of tobacco streak virus on Lablab purpureus. New Dis Rep 28:11 Briddon RW (2001) Begomovirus. Geminiviridae. The Springer index of viruses. Springer, New York, pp 340–348, https://doi.org/10.1007/3-540-31042-8_55. Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Capoor SP, Varma PM (1950) A new virus disease of Dolichos lablab. Curr Sci 19:248–249 Kiranmai G, Sreenivasulu P, Nayudu MV (1998) Epidemiology of cucumber mosaic cucumovirus isolates naturally infecting three solanaceous vegetable crops around Tirupati. Indian Phytopathol 51:315–318 Li CS (1991) Identification of clover yellow vein virus infecting Dolichos lablab. Virol Sin 6:223–226 Maruthi MN, Rekha AR, Govindappa MR, Colvin J, Muniyappa V (2006a) A distinct Begomovirus causes Indian dolichos yellow mosaic disease. Plant Pathol 55:290–294 Maruthi MN, Manjunatha B, Rekha AR, Govindappa MR, Colvin J, Muniyappa V (2006b) Dolichos yellow mosaic virus belongs to a distinct lineage of old world begomoviruses; its biological and molecular properties. Ann Appl Biol 149:187–195

926

Dracaena spp. (Cornstalk dracaena)

Odedara OO, Hughes J’A, Odebode AC, Odu BO (2008) Multiple virus infections of lablab [Lablab purpureus (L.) Sweet] in Nigeria. J Gen Plant Pathol 74:322–325 Pavithra BS, Krishna Reddy M, Jalali S, Samuel DK (2017) Molecular detection of bean common mosaic necrosis virus in Dolichos bean (Lablab purpureus). p 118. In: 26th Annual Conference of Indian Virological Society, VIROCON 2017 (Abstract). Pudashini BJ, Shahid MS, Natsuaki KT (2013) First report of bean common mosaic necrosis virus (BCMNV) infecting sweet bean in Nepal. Plant Dis 97:290 Qazi J, Ilyas M, Mansoor S, Briddon RW (2007) Legume yellow mosaic viruses: genetically isolated begomoviruses. Mol Plant Pathol 8(4):343–348 Raj SK, Aslam N, Srivastava KM, Singh BP (1988) Association of geminivirus like particles with yellow mosaic disease of Dolichos lablab. Curr Sci 58:813–814 Rishi N, Dhawan P (1987) Bean mosaic and bean yellow mosaic diseases of Hyacinth bean (Lablab purpureus L.) sweet and screening of available genotypes to find the sources of resistance. Indian J Pl Pathol 5:63–68 Sengooba TN, Spence NJ, Walkey DGA, Allen DJ, Femi Lana A (1997) The occurrence of bean common mosaic necrosis virus in wild and forage legumes in Uganda. Plant Pathol 46:95–103 Singh PK, Rai N, Singh DV, Singh AP (2012) Incidence of Dolichos yellow mosaic virus (DYMV) and yield potential in Indian bean (Lablal purpureus) F1’S. J Agric Technol 8:1469–1474 Subramanian KS (1975) Studies on yellow mosaic disease of Lablab niger Medikus (Dolichos lablab L.). Ph.D. Thesis, Tamil Nadu Agricultural University, Coimbatore, 102 p. Udayashankar AC, Chandra Nayaka S, Niranjana SR, Lund OS, Prakash HS (2011) First report of bean common mosaic virus infecting Lablab purpureus in India. Plant Dis 95:881 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133 Zhou GY, Chen YX (1990) Identification of staple viruses of dolichos in Nanjing. Chin J Virol 6:233–238

Dracaena spp. (Cornstalk dracaena) Family: Asparagaceae

Dracaena mottle virus

Ornamental

(DrMV)

Taxonomic position DrMV is a tentative member of the genus Badnavirus and family Caulimoviridae. Geographical distribution DrMV-infected plants of Dracaena spp. were reported from New Zealand, China, and the USA (Su et al. 2007; Baker and Jeyaprakash 2014). Symptoms and host(s) The virus-infected dracaena plants exhibit mottling and chlorotic patches on the leaves. Dracaena spp. is the only natural hosts of this virus. Transmission The virus is mechanically sap-transmissible. Virion properties and genome The virions are bacilliform-like particles approximately 125–130 nm long and 30 nm wide. The genome is a double-stranded DNA of 7531 bp (NC_008034) and possesses seven putative ORFs on

Dracaena spp. (Cornstalk dracaena)

927

the plus strand that potentially encodes proteins of 17.6, 14.9, 215.0, 11.9, 11.3, 16.1, and 11.0 kD (Su et al. 2007).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Tospovirus

(INSV)

Family: Peribunyaviridae

D

INSV infection in plants of Dracaena fragrans was reported from Iran (Ghotbi and Shahraeen 2012; Ghotbi 2013). The virus-infected dracaena plants exhibit symptoms of necrotic leaf spot, chlorosis, and stunting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Tospovirus

(TSWV)

Family: Peribunyaviridae

TSWV infection has been reported in plants of Dracaena spp. from Iran and the USA (Hausbeck et al. 1992; Ghotbi and Shahraeen 2012). The virus-infected dracaena plants exhibit mottle mosaic, necrosis, and chlorotic spotting symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow ring virus

(TYRV)

Taxonomic position TYRV is a tentative member of the genus Tospovirus and family Peribunyaviridae TYRV was detected in plants of Dracaena fragrans in Iran (Ghotbi and Shahraeen 2012). The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of TYRV, refer to Solanum lycopersicum.

References Baker CA, Jeyaprakash A (2014) Dracaena mottle virus in lucky bamboo. Plant Pathology Circular No 413, Fla Dept Agric and Consumer Serv Division of Plant Industry, Florida, USA. Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iran J Plant Pathol 49:41 Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses affecting propagated ornamentals in Northern Iran. Int Res J Microbiol 3:373–381 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among green house ornamentals in Pennsylvania. Plant Dis 76:795–800 Su L, Goa S, Juang Y, Chaogun J, Wang S, Ma Y, Fang R, Chen X (2007) Complete genomic sequence of Dracaena mottle virus, a distinct badnavirus. Virus Genes 35:423–429

928

Drakaea spp. (Drakaea elastica, Drakaea livida) (Hammer orchids)

Drakaea spp. (Drakaea elastica, Drakaea livida) (Hammer orchids) Family: Orchidaceae

Ornamental

Caladenia virus A

(CalVA)

Taxonomic position Genus: Poacevirus

Family: Potyviridae

CalVA infection in plants of Drakaea elastica was reported from Australia (Wylie et al. 2012). The virus is possibly transmitted mites. For more details of CalVA, refer to Caladenia spp.

Donkey orchid symptomless virus Taxonomic position Genus: Platypuvirus

(DOSV)

Family: Alphaflexiviridae

DOSV infection in plants of Drakaea elastica was reported from Western Australia (Ong et al. 2016). The virus-infected hammer orchid plants do not exhibit any symptoms. The virus is mechanically saptransmissible. For more details of DOSV, refer to Diuris spp.

Drakaea virus A Taxonomic position Genus: Goravirus

(DVA)

Family: Virgaviridae

Geographical distribution DVA infection in plants of Drakaea livida was reported from Australia (Ong et al. 2016). Symptoms and host(s) The virus-infected hammer orchid plants exhibit slight leaf discoloration symptoms. Transmission No vector is known for this virus. The virus is mechanically sap-transmissible. Virion properties and genome The virions are rod-shaped and about 20nm in width, but it was difficult to determine their length because of disintegration during purification. The genome has two single-stranded RNAs of 4490 nt (RNA 1; KP760461) and 2905 nt (RNA 2; KP760462). There is a 30 -terminal tRNA-like structure on both genomic RNAs (Adams et al. 2017).

Duranta spp.

929

References Adams MJ, Adkins S, Bragard C, Gilmer D, Li D, MacFarlane SA, Wong S-M, Melcher U, Ratti C, Ryu KH, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Virgaviridae. J Gen Virol 98:1999–2000 Ong JWL, Phillips RD, Dixon KW, Jones MGK, Wylie SJ (2016) Characterization of the first two viruses described from wild populations of hammer orchids (Drakaea spp.) in Australia. Plant Pathol 65:163–172 Wylie SJ, Tan AJY, Li H, Dixon KW, Jones MGK (2012) Caladenia virus A, an unusual new member of the family Potyviridae from terrestrial orchids in Western Australia. Arch Virol 157:2447–2452

D Duboisia spp. Family: Solanaceae

Medicinal

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Tospovirus

(TSWV)

Family: Peribunyaviridae

TSWV infection in plants of Duboisia spp. was reported from Queensland (Greber and McCarthy 1977; McCarthy and Greber 1978). The virus-infected Duboisia plants exhibit symptoms of leaf veinbanding, line patterns, dieback, leaf fall, and wilting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Greber RS, McCarthy GJP (1977) A disease of Duboisia caused by tomato spotted wilt virus (TSWV). Aust Plant Pathol Soc Newsl 6:54–55 McCarthy GJP, Greber RS (1978) Tomato spotted wilt virus in Duboisia. Queensland Agric J 104:336–338

Duranta spp. Family: Verbenaceae

Ornamental

Catharanthus yellow mosaic virus Taxonomic position Genus: Begomovirus

(CatYMV)

Family: Geminiviridae

CatYMV infection in plants of Duranta repens was reported from Pakistan (Mustujab et al. 2015). The virus-infected duranta plants exhibit mild leaf curl symptoms. The virus is transmitted by the whitefly

930

Duranta spp.

vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of CatYMV, refer to Catharanthus spp.

Clerodendron yellow mosaic virus Taxonomic position Genus: Begomovirus

(ClYMV)

Family: Geminiviridae

ClYMV infection in plants of Duranta erecta was reported from India (Jaidi et al. 2015). The virusinfected duranta plants show severe yellow mosaic with upward leaf curling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ClYMV, refer to Clerodendrum spp.

Duranta leaf curl virus Taxonomic position Genus: Begomovirus

(DuLCV)

Family: Geminiviridae

Geographical distribution DuLCV infection in plants of Duranta spp. was reported from Pakistan and India (Iram et al. 2005; Tahir et al. 2006; Sharma et al. 2007; Marwal et al. 2013; Anwar and Tahir 2018). Symptoms and host(s) The virus-infected duranta plants exhibit symptoms of twisting, upward leaf curling, chlorosis, and reduction of leaves (Iram et al. 2005; Sharma et al. 2007). Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome contains a single molecule of circular single-stranded DNA (DNA-A) of 2759 nt (KT948069) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2018; Anwar and Tahir 2018).

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Duranta repens was reported from Pakistan (Tahir et al. 2006). The virus-infected duranta plants exhibit leaf curl symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner, and also by grafting. For more details of ToLCNDV, refer to Solanum lycopersicum.

Duranta spp.

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References Anwar S, Tahir M (2018) Identification of a new begomovirus infecting Duranta repens in Pakistan. Arch Virol 163:809–813 Briddon RW (2001) Begomovirus. Geminiviridae. The Springer index of plant viruses. Springer, New York, pp 340-348, https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A. (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Iram S, Amrao L, Mansoor MS, Malik AH, Briddon RW, Zafar Y (2005) First report of a begomovirus associated with leaf curl disease of Duranta erecta in Pakistan. Plant Pathol 54:260 Jaidi M, Kumar S, Srivastava A, Raj SK (2015) First report of Clerondendron yellow mosaic virus on golden dewdrop (Duranta erecta) in India. New Dis Rep 32:12 Marwal A, Sahu AK, Gaur RK (2013) Molecular characterization of a begomovirus infecting a new host golden duranta (Duranta erecta) in India. Int J Curr Microbial Appl Sci 2:45–48 Mustujab A, Briddon RW, Tahir M (2015) First report of Catharanthus yellow mosaic virus infecting Duranta repens in Pakistan. J Plant Pathol 97:210 Sharma A, Shankarappa KS, Rangaswamy KT, Dubey RK, Maruthi MN (2007) Detection and characterization of a Begomovirus associated with leaf curl disease of Duranta (Duranta erecta). In: Proceedings of the international satellite symposium on viruses of ornamental and temperate fruit crops, IHBT, Palampur, India, December 17–18, 2007, p 8 Tahir M, Haider MS, Shah AH, Rashid N, Saleem F (2006) First report of bipartite begomovirus associated with leaf curl disease of Duranta repens in Pakistan. J Plant Pathol 88:339 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

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Ecballium elaterium (Squirting cucumber) Family: Cucurbitaceae

Medicinal

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

CGMMV infection in plants of Ecballium elaterium was reported from Israel (Antignus et al. 1990; Shargil et al. 2017). The virus-infected squirting cucumber plants exhibit yellow green spots and vein-clearing on young leaves and necrosis at later stages of growth. The virus is not transmitted by any insect vector. The virus is transmissible through soil and irrigation water contaminated with infected plant debris. The virus is transmissible by mechanical sap-inoculation, and grafting. The virus is also pollen transmitted. For more details of CGMMV, refer to Cucumis sativus.

Cucumber vein yellowing virus Taxonomic position Genus: Ipomovirus

(CVYV)

Family: Potyviridae

CVYV infection in plants of Ecballium elaterium was reported from Spain (Janssen et al. 2002). The virus-infected squirting cucumber plants exhibit vein-clearing and leaf chlorosis symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CVYV, refer to Cucumis sativus.

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(CABYV)

Family: Luteoviridae

© Springer Nature India Private Limited 2019 K. S. Sastry et al., Encyclopedia of Plant Viruses and Viroids, https://doi.org/10.1007/978-81-322-3912-3

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Echinacea purpurea (Purple coneflower)

CABYV infection in plants of Ecballium elaterium was reported from Tunisia (Hattab et al. 2005, 2009). The virus-infected squirting cucumber plants exhibit severe yellowing symptoms on older leaves. The virus is transmitted by aphids, Myzus persicae, M. euphorbiae, and Aphis gossypii, in a persistent, circulative, non-propagative manner (Lecoq et al. 1992). The virus is not transmissible by mechanical sap-inoculation. For more details of CABYV, refer to Cucumis melo.

Zucchini yellow fleck virus Taxonomic position Genus: Potyvirus

(ZYFV)

Family: Potyviridae

ZYFV infection in plants of Ecballium elaterium was reported from Israel, Greece, and France (Avgelis 1985; Gilbert-Albertini and Lecoq 1994; Antignus et al. 1995; Desbiez et al. 2007). The virus-infected squirting cucumber plants exhibit symptoms of mosaic, chlorotic spots, and fruit deformations. The virus is transmitted by the aphid vector, Myzus persicae, in a non-persistent manner, and is also transmissible through mechanical sap-inoculation. For more details of ZYFV, refer to Cucurbita pepo.

References Antignus Y, Pearlsman M, Ben-Yoseph R, Cohen S (1990) Occurrence of variant of Cucumber green mottle mosaic virus in Israel. Phytoparasitica 18:50–56 Antignus Y, Levy D, Cohen S (1995) Characterisation of a variant of zucchini yellow fleck virus (ZYFV), a potyvirus causing a wilt disease of melons in Israel. Ann Appl Biol 126(1):111–120 Avgelis AD (1985) Epidemiological studies of Zucchini yellow fleck virus in Crete. Phytopath Mediterr 24:208–210 Desbiez C, Justafre I, Lecoq H (2007) Molecular evidence that Zucchini yellow fleck virus is a distinct and variable potyvirus related to Papaya ring spot virus and Moroccan watermelon mosaic virus. Arch Virol 152:449–455 Gilbert-Albertini F, Lecoq H (1994) The characterization of a strain of Zucchini yellow fleck virus found in southeastern France. J Phytopathol 140:375–384 Hattab MM, Kummert J, Roussel S, Ezzaier K, Zouba A, Jijakli MH (2005) First report of Cucurbit aphid-borne yellows virus in Tunisia causing yellows on five cucurbitaceous species. Plant Dis 89:776 Hattab MM, Guathier N, Zouba A (2009) Biological and molecular characterization of the Cucurbit aphid-borne yellows virus affecting cucurbits in Tunisia. Plant Dis 93:1065–1072 Janssen D, Ruiz L, Velasco L, Segundo E, Cuadrado IM (2002) Non-cucurbitaceous weed species shown to be natural hosts of Cucumber vein yellowing virus in southeastern Spain. Plant Pathol 51:797 Lecoq H, Bourdin D, Wipf-Scheibel C, Bon M, Lot H, Lemaire O, Herrbach E (1992) A new yellowing disease of cucurbits caused by a Luteovirus, Cucurbit aphid-borne yellows virus. Plant Pathol 41:749–761 Shargil D, Smith E, Lachman O, Reingold V, Darzi E, Tam Y, Dombrovsky A (2017) New weed hosts for Cucumber green mottle mosaic virus in wild Mediterranean vegetation. Eur J Plant Pathol 148:473–480

Echinacea purpurea (Purple coneflower) Family: Asteraceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Medicinal

(AMV)

Family: Bromoviridae

Echinacea purpurea (Purple coneflower)

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AMV infection in plants of Echinacea purpurea was reported from Bulgaria (Dikova et al. 2010; Dunich and Mishchenko 2015). The virus-infected purple coneflower plants exhibit spotting and dwarfing symptoms. The virus is transmitted by several species of aphids in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Broad bean wilt virus 2

(BBWV-2)

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Taxonomic position Genus: Fabavirus

Family: Secoviridae

BBWV-2 infection in plants of Echinacea purpurea was reported from China (Li et al. 2012; Dunich and Mishchenko 2015). The virus-infected purple coneflower plants exhibit symptoms of necrosis, leaf rolling, yellow mosaic, and mosaic in leaves. The virus is transmitted by the aphid vector, Acyrthosiphon solani, in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BBWV-2, refer to Vicia faba.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Echinacea purpurea was reported from the USA, Hungary, Bulgaria, China, Italy, Germany, New Zealand, and Japan (EU432180) (Muehle and Schumann 1964; Yamamoto et al. 1993; Horvath et al. 2006; Guifen et al. 2007; Dikova et al. 2010; Davino et al. 2012; Dunich and Mishchenko 2015). The virus-infected purple coneflower plants exhibit symptoms of mosaic, yellow mosaic, variegation on the leaves, yellow mottling in leaves which were often malformed, with ringspots, and leaf deformation. The virus is transmitted by the aphid vector, Aphis gossypii, in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Potato virus Y

(PVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

PVY infection in plants of Echinacea purpurea was reported from Bulgaria, Hungary, and Ukraine (Horvath et al. 2006; Dashchenko 2014; Dunich and Mishchenko 2015). The virus-infected purple coneflower plants exhibit leaf curling and deformation symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts and by grafting. For more details of PVY, refer to Solanum tuberosum.

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Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

Echinacea purpurea (Purple coneflower)

(TMV)

Family: Virgaviridae

TMV infection in plants of Echinacea purpurea was reported from Hungary and Bulgaria (Horvath et al. 2006; Dikova et al. 2013; Dunich and Mishchenko 2015). The virus-infected purple coneflower plants exhibit mosaic and spotting symptoms. There is no known vector for this virus. The virus is mechanically sap-transmissible and is also transmissible through contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in plants of Echinacea purpurea was reported from Lithuania (Samuitien_e and Navalinskien_e 2010; Dunich and Mishchenko 2015). The virus-infected purple coneflower plants exhibit stunting, leaf malformation, various shaped chlorotic spots, and ringspot symptoms. The virus is transmitted by nematode vectors and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Echinacea purpurea was reported from Lithuania (Samuitien_e and Navalinskien_e 2010; Dunich and Mishchenko 2015). The virus-infected purple coneflower plants exhibit symptoms of stunting, leaf malformation, various shaped chlorotic spots, and ringspots. The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Echinacea purpurea was reported from Lithuania (Dunich and Mishchenko 2015). The virus-infected purple coneflower plants exhibit symptoms of stunting, leaf malformation, various shaped chlorotic spots, and ringspots. The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation, and by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Echinochloa crus-galli (Cockspur grass)

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Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Echinacea purpurea was reported from Europe and Hungary (Horvath et al. 2006; Dikova 2011; Dunich and Mishchenko 2015). The virus-infected purple coneflower plants exhibit symptoms of yellow spotting on the leaves, chlorotic or dark red ringspots turning into brown necrotic lesions. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Dashchenko AV (2014) Monitoring of viruses of medicinal plants of the family Asteraceae. Karantyn I Zahyst Roslyn 1:10–14 Davino S, Panno S, Rangel EA, Davino M, Bellardi MG, Rubio L (2012) Population genetics of Cucumber mosaic virus infecting medicinal, aromatic and ornamental plants from northern Italy. Arch Virol 157:739–745 Dikova B (2011) Tomato spotted wilt virus on some medicinal and essential oil-bearing plants in Bulgaria. Bulg J Agric Sci 17:306–313 Dikova B, Djourmanski A, Lambev H (2010) Isolation of viruses (Polyphagues) from some important medicinal plants in Bulgaria. J Balkan Ecology 13:33–45 Dikova B, Djourmanski A, Lambev H (2013) Establishment of economically important viruses on Echinacea purpurea and their influence on yield. In: Pospelov S (ed) Proceedings of the conference “Innovative aspects to coneflower studies”. Dyvosvit, Polteva, pp 36–45 Dunich AA, Mishchenko LT (2015) Purple coneflower viruses: species diversity and harmfulness. Biopolym Cell 31(1):15–28 Guifen L, Shuifang Z, Qun Z (2007) Identification of Cucumber mosaic virus isolated from Echinacea purpurea. Plant Protect Beijing 33:54–56 Horvath J, Baracsi E, Takacs A, Kazinczi G, Gaborjanyi R, Krajczinger R (2006) Virus infection of ornamental plants in Hungary. Cereal Res Commun 34:485–488 Li GF, Wei MS, Ma J, Zhu SF (2012) First report of Broad bean wilt virus 2 in Echinacea purpurea in China. Plant Dis 96:1232 Muehle E, Schumann K (1964) On the presence of Cucumber mosaic virus (Marmor cucumeris H.) on Echinacea purpurea (L.) Moench. Pharmazie 19:417–421 Samuitien_e M, Navalinskien_e M (2010) Association of tobacco rattle and tobacco ringspot viruses with purple coneflower disease. Bot Lithuanica 16:51–56 Yamamoto T, Ishii M, Sasaya T, Iwasaki M (1993) Mosaic disease of Echinacea (Echinacea purpurea, Compositae). Proc Assoc Plant Protect Shikoku 28:49–53

Echinochloa crus-galli (Cockspur grass) Family: Poaceae

Forage crop

Barley yellow dwarf virus PAV Genus: Luteovirus

(BYDV-PAV)

Family: Luteoviridae

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Echinochloa crus-galli (Cockspur grass)

BYDV-PAV infection in plants of Echinochloa crusgalli was reported from Syria (Ansi et al. 2007). The virus is transmitted by a aphid vectors in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. For more details of BYDV-PAV, refer to Hordeum vulgare.

Echinochloa hoja blanca tenuivirus Taxonomic position Genus: Tenuivirus

(EHBV)

Family: Phenuiviridae

Geographical distribution EHBV infection in plants of Echinochloa crus-galli was reported from Latin America (de Miranda et al. 1996a, b, c). Symptoms and host(s) The virus-infected cockspur grass plants exhibit striping or mottling symptoms on the leaves. The virus infects plants of the family Poaceae. Transmission The virus is transmitted by the planthopper vector, Tagosodes cubanus, in a circulative, propagative manner (Falk and Tsai 1998). The virus is not mechanically sap-transmissible. Virion properties and genome The ribonucleoproteins (RNPs) have a thin filaments shape; they consist of nucleocapsids, 3–10 nm in diameter, with lengths proportional to the sizes of the RNAs they contain. The filaments particles may appear to be spiral-shaped, branched, or circular. No envelope has been observed. The genome is segmented and consists of five or more segments of linear negative-sense and ambisense single-stranded RNA. Three of the genome segments have been sequenced: RNA3 contains 2336 nt (L75930), RNA4 of 1874 nt (L48441), and RNA5 of 1334 nt (L47430) (de Miranda et al. 1996a, b, c; Falk and Tsai 1998).

Echinochloa ragged stunt virus Taxonomic position Genus: Oryzavirus

(ERSV)

Family: Reoviridae

Geographical distribution ERSV was first reported in plants of Echinochloa crus-galli var. oryzicola from Taiwan by Chen et al. (1980). The virus spreads in Taiwan (Chen et al. 1986). Symptoms and host(s) The virus-infected cockspur grass plants exhibit symptoms of tillering, severe dwarfing, malformation of leaves with serrated margins, and veinal swellings. Plants appear greener and fail to head or show a delay in heading time leading to empty grains. The virus naturally infects plants in the family Poaceae, including Echinochloa crus-galli var. oryzicola, Setaria italica, and Triticum aestivum (wheat); additional species are experimental hosts.

Echinochloa crus-galli (Cockspur grass)

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Transmission The virus is transmitted by planthopper vectors, Sogatella longifurcifera and S. vibix in a persistent manner. The minimum acquisition access period for the vector is about 3 h, the latent period is about 9 days, and the minimum inoculation access time is about 1 h. The virus is not passed through the egg. The virus is not transmissible by mechanical inoculation, by contact between plants, by seed, nor by pollen.

Virion properties and genome The virions are non-enveloped, icosahedral with a double capsid structure, and about 70 nm in diameter. Pentameric turrets (VP2) sit on the outside of the innermost capsid (VP3). Each turret is surrounded by five peripheral trimers (VP9), and each trimer binds to three clamp proteins (VP4B). The outer incomplete capsid has a T = 13 icosahedral symmetry and the inner capsid a T = 2 icosahedral symmetry. The genome has ten segments of linear double-stranded RNA coding for 12 proteins (Francki and Boccardo 1983; Upadhyaya and Waterhouse 2011). No sequences are currently available.

Wheat streak mosaic virus Taxonomic position Genus: Tritimovirus

(WSMV)

Family: Potyviridae

WSMV infection in plants of Echinochloa crus-galli was reported from Kansas, USA (Christian and Willis 1993). The virus is transmitted by eriophyid mite vectors in a semi-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of WSMV, refer to Triticum aestivum.

References Ansi A, Kumari SG, Haj Kasem A, Makkouk KM, Muharram I (2007) The occurrence of Barley yellow dwarf viruses on cereal crops and wild grasses in Syria. Arab J Pl Prot 25:1–9 Chen CC, Chiu RJ, Chen MJ, Wang YS, Ko WH (1980) Plant Prot Bull (Taiwan) 22:425 Chen CC, Chen MJ, Chiu RJ (1986) Echinochloa ragged stunt: symptomatology, host range and transmission. Plant Prot Bull (Taiwan) 28:371–381 Christian ML and Willis WG (1993) Survival of wheat streak mosaic virus in grass hosts in Kansas for wheat harvest to fall wheat emergence. Plant Dis 77:239–242 Falk BW, Tsai JH (1998) Biology and molecular biology of viruses in the genus Tenuivirus. Annu Rev Phytopathol 36:139–163 Francki RIB, Boccardo G (1983) The plant Reoviridae. In: Joklik WK (ed) The Reoviridae. Plenum Press, New York, pp 505–563 de Miranda JR, Munoz M, Madriz J, Wu R, Espinoza AM (1996a) Sequence of Echinochloa hoja blanca tenuivirus RNA3. Virus Genes 13:65–68 de Miranda JR, Munoz M, Wu R, Espinoza AM (1996b) Sequence of Echinochloa hoja blanca tenuivirus RNA-4. Virus Genes 13:61–64 de Miranda JR, Munoz M, Wu R, Espinoza AM (1996c) Sequence of Echinochloa hoja blanca tenuivirus RNA-5. Virus Genes 12:131–134 Upadhyaya NM, Waterhouse PM (2011) Oryzavirus. Reoviridae. In: The Springer index of viruses. Springer, New York, pp 1621–1626. https://doi.org/10.1007/978-0-387-95919-1_265

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Echium candicans (Pride of Madeira)

Echium candicans (Pride of Madeira) Synonyms Echium fastuosum Family: Boraginaceae

Ornamental

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Echium candicans was reported from France (Cardin and Moury 2007). The virus-infected Pride of Madeira plants exhibit mosaic symptoms. The virus is transmitted by a number of aphids in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of host plants. For more details of CMV, refer to Cucumis sativus.

Reference Cardin L, Moury B (2007) First report of cucumber mosaic virus in Echium candicans in France. Plant Dis 91:1516

Eclipta prostrata (False daisy) Synonyms Eclipta alba Family: Asteraceae

Medicinal

Alternanthera yellow vein virus Taxonomic position Genus: Begomovirus

(AlYVV)

Family: Geminiviridae

AlYVV infection in plants of Eclipta prostrata was reported from Guangzhou, Guangdong province (China), and Pakistan (He et al. 2008; Paul et al. 2008; Zhang et al. 2015; Zaidi et al. 2017). The virusinfected false daisy plants exhibit yellow vein symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. For more details of AlYVV, refer to Alternanthera spp.

Eclipta prostrata (False daisy)

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Eclipta yellow vein virus Taxonomic position Genus: Begomovirus

(EYVV)

Family: Geminiviridae

Geographical distribution EYVV infection in plants of Eclipta prostrata was reported from Pakistan (Khatri et al. unpublished – GQ478343). Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2740 nt (GQ478343) (Brown et al. 2015; Zerbini et al. 2017).

Papaya leaf curl China virus Taxonomic position Genus: Begomovirus

(PaLCuCNV)

Family: Geminiviridae

PaLCuCNV infection in plants of Eclipta prostrata was reported from China (Zhang et al. 2015). The virus-infected false daisy plants exhibit vein yellowing symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. For more details of PaLCuCNV, refer to Carica papaya.

Pepper yellow vein Mali virus Taxonomic position Genus: Begomovirus

(PepYVMLV)

Family: Geminiviridae

PepYVMLV infection in plants of Eclipta prostrata was reported from China (Zhang et al. 2015). The virus-infected false daisy plants exhibit vein yellowing symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. For more details of PepYVMLV, refer to Capsicum annuum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Eclipta prostrata was reported from Australia (Sharman et al. 2015). The virus-infected false daisy plants exhibit leaf deformation symptoms. The virus is transmitted by the thrips vectors, the virus present in/on pollen, entering the host though injuries caused by thrips while

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Eclipta prostrata (False daisy)

feeding. The virus is also mechanically sap-transmissible but is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Eclipta prostrata was reported from Pakistan (Haider and Tahir 2005; Haider et al. 2006). The virus-infected false daisy plants exhibit yellow vein symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner, and also by grafting. For more details of ToLCNDV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Eclipta prostrata was reported from Georgia (USA) (Mullis et al. 2009). The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Haider MS, Tahir M (2005) A new species of Begomovirus infecting Capsicum annuum grossum (Bell Pepper) and prevalence of Tomato Leaf Curl New Delhi Virus in Momordica charantia and Eclipta prostrata under natural conditions of Pakistan. 2nd Joint Conference of the International Working Groups on Legume (IWGLV) and Vegetable Viruses (IWGVV), April 10–14 (2005), held at Florida, USA Haider MS, Tahir M, Latif S, Briddon RW (2006) First report of Tomato leaf curl New Delhi virus infecting Eclipta prostrata in Pakistan. Plant Pathol 55:285 He ZF, Mao MJ, Yu H, Wang XM, Li HP (2008) First report of a strain of Alternanthera yellow vein virus infecting Eclipta prostrata (L.) L. (Compositae) in China. J Phytopathol 156:496–498 Mullis S, Bertrand PF, Brown SL, Csinos A, Diaz-Perez JC, Gitaitis R, Hickman LL, Johnson A, LaHue SS, Martinez N, McPherson RM, Nischwitz C, Reay-Jones F, Riley D, Sherwood J, Stevenson K, Wells L (2009) Tospoviruses in Solanaceae and other crops in the coastal plain of Georgia. University of Georgia, College of Agricultural and Environmental Sciences, Bulletin 1354, pp 51 Paul S, Ghosh R, Das S, Acharyya S, Palit P, Das A, Mir JI, Ghosh SK, Roy A (2008) Detection of a Begomovirus associated with yellow vein disease of Eclipta prostrata in eastern India. Indian J Virol 19:120 Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207 Zaidi SS, Shakir S, Farooq M, Amin I, Mansoor S (2017) First report of Alternanthera yellow vein virus from Eclipta prostrata in Pakistan. Plant Dis 101:266 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zhang J, Jia SP, Yang CX, Liu Z, Wu ZJ (2015) Detection and molecular characterization of three Begomoviruses associated with yellow vein disease of Eclipta Prostrata in Fujian, China. J Plant Pathol 97:161–165

Elaeis guineensis (Oil palm)

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Edgeworthia chrysantha (Oriental paperbush) Family: Thymelaeaceae

Commercial crop

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Edgeworthia chrysantha was reported from France and Italy (Cardin and Moury 2009). The virus-infected oriental paperbush plants exhibit mosaic and oak leaf pattern symptoms. The virus is transmitted by the aphid vector, Myzus persicae, in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Reference Cardin L, Moury B (2009) First report of Cucumber mosaic virus in Edgeworthia chrysantha in France and Italy. J Plant Pathol 91(2):502

Elaeis guineensis (Oil palm) Family: Arecaceae

Oil seed

African oil palm ringspot virus Taxonomic position Genus: Robigovirus

(AOPRV)

Family: Betaflexiviridae

Geographical distribution AOPRV infection in plants of Elaeis guineensis was reported from South America (Morales et al. 2002; Lozano et al. 2010). Symptoms and host(s) The virus-infected oil palm plants exhibit lethal ringspot symptoms. Transmission The virus is mechanically sap-transmissible. There is no known vector for this virus. Virion properties and genome The virions are flexuous filaments 725–825 nm long and 12 nm wide. The genome consists of a single molecule of positive-sense, single-stranded RNA of 7781 nt with five ORFs: a large (209 kDa)

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Elaeis guineensis (Oil palm)

replication protein, a triple gene block (associated with cell-to-cell movement), and a single 29 kDa coat protein (AY072921) (Morales et al. 2002).

Coconut cadang-cadang viroid Taxonomic position Genus: Cocadviroid

(CCCVd)

Family: Pospiviroidae

CCCVd infection in plants of Elaeis guineensis was reported from the Philippines, Southeast Asia, South Pacific, and Malaysia (Randles et al. 1980, 1998; Hanold and Randles 1991; Vadamalai et al. 2006, 2008; Wu et al. 2013; Roslan et al. 2016). Previously CCCVd was described as “genetic” orange spotting (GOS) (Smilde 1962; Forde and Leyritz 1968; Robertson et al. 1968; Turner 1981). The viroid-infected African oil palm leaves develop orange color spots on pinnae of older fronds, and the youngest opened fronds became chlorotic and developed smaller spots. There is premature loss of male florets in the early stage of infection. Diseased trees did not produce inflorescences and showed broken pinnae, and the number and size of fronds were reduced (Randles et al. 1980). The viroid is mechanically sap-transmissible and is also transmissible through contaminated implements. For more details of CCCVd, refer to Cocus nucifera.

References Forde SCM, Leyritz MJP (1968) A study of confluent orange spotting of the oil palm in Nigeria. J Niger Inst Oil Palm Res 4:371–380 Hanold D, Randles JW (1991) Detection of coconut cadang cadang viroid-like sequences in oil and coconut palm and other monocotyledons in the south-west Pacific. Ann Appl Biol 118:139–151 Lozano I, Morales FJ, Martinez AK, Pean EA (2010) Molecular characterization and detection of African oil palm ring spot virus. J Phytopathol 158:167–172 Morales FJ, Lozano I, Velasco AC, Arroyave JA (2002) Detection of a fovea-like virus in African oil palms affected by a lethal ringspot disease in South America. J Phytopathol 150:611–615 Randles JW, Boccardo G, Imperial JS (1980) Detection of the cadang-cadang associated RNA in African oil palm and buri palm. Phytopathology 70(3):185–189 Randles JW, Hanold D, Pacumbaba EP, Rodriguez MJB (1998) Cadang-cadang disease of coconut palm – an overview. In: Randles JW, Hanold D (eds). Report on ACIAR-funded research on viroids and viruses of coconut palm and other tropical monocotyledons 1985–1993. ACIAR Working Paper No. 51 August 1998. ISBN 1 86320 219 6, Australian Centre for International Agricultural Research, Canberra, Australia. Robertson JS, Prendergast AG, Sly JMA (1968) Diseases and disorders of the oil palm (Elaeis guineensis) in West Africa. J Niger Inst Oil Palm Res 4:381–409 Roslan ND, Meilina OA, Mohamed-Azni I-NA, Seman IA, Sundram S (2016) Comparison of RNA extraction methods for RT-PCR detection of Coconut cadang-cadang viroid variant in orange spotting oil palm leaves. Can J Plant Pathol 38:382–388 Smilde KW (1962) 10th Annual Report of the West African Institute for Oil Palm Research. pp 71–78 Turner PD (1981) Oil palm diseases and disorders. Oxford University Press, Kuala Lumpur Vadamalai G, Hanold D, Rezaian MA, Randles JW (2006) Variants of Coconut cadang-cadang viroid isolated from an African oil palm (Elaies guineensis Jacq.) in Malaysia. Arch Virol 151:1447–1456 Vadamalai G, Perera AAFLK, Hanold D, Rezaian MA, Randles JW (2008) Detection of Coconut cadang-cadang viroid sequences in oil palm and coconut palm by ribonuclease protection assay. Ann Appl Biol 154:117–125 Wu YH, Cheong LC, Meon S, Lau WH, Kong LL, Joseph H, Vadamalai G (2013) Characterization of coconut cadangcadang viroid variants from oil palm affected by orange spotting disease in Malaysia. Arch Virol 158:1407–1410

Elettaria cardamomum (Small cardamom; Queen of spices)

945

Eleocharis dulcis (Water chestnut) Family: Cyperaceae

Tuber crop

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Eleocharis dulcis was reported from China (Liu et al. 2014). The virusinfected water chestnut plants exhibit mosaic, chlorotic, dwarfing, and malformations. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of CMV, refer to Cucumis sativus.

Reference Liu J, Wang YF, Hong N, Wang GP, Wang LP (2014) The first report of Cucumber mosaic virus infecting water chestnut in China. Plant Dis 98:164

Elettaria cardamomum (Small cardamom; Queen of spices) Family: Zingiberaceae

Spice crop

Banana bract mosaic virus Taxonomic position Genus: Potyvirus

(BBrMV)

Family: Potyviridae

BBrMV infection in plants of Elettaria cardamomum was reported from Kerala, Karnataka, and Tamil Nadu in India (Siljo et al. 2011, 2012, 2014; Siljo and Bhat 2014). The virus-infected small cardamom plants exhibit symptoms of continuous or discontinuous spindle-shaped yellow or light green intravenous streaks along the veins and midrib. These streaks later coalesce together and impart a yellow or light green color to the veins. Discontinuous spindle-shaped mottling on the pseudostem and petioles was also noticed. In severe cases, tillering in an infected plant was suppressed (Siljo et al. 2012). The virus is transmitted primarily through infected suckers, and secondary spread is by aphids in a nonpersistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BBrMV, refer to Musa spp.

Cardamom mosaic virus Synonyms Indian cardamon mosaic virus

(CdMV)

E

946

Taxonomic position Genus: Macluravirus

Elettaria cardamomum (Small cardamom; Queen of spices)

Family: Potyviridae

Geographical distribution CdMV infection in plants of Elettaria cardamomum was first reported from South India (Uppal et al. 1945). The virus spreads in Guatemala, Sri Lanka, the USA, and India (Capoor 1967; Rao and Naidu 1973; Rao 1977; Dimitman et al. 1984; Gonsalves et al. 1986; Saigopal et al. 1990; Venugopal 1995; Jacob et al. 2003; Biju et al. 2010; Siljo et al. 2014). This virus disease was earlier described in India as “Katte” or marble disease. Symptoms and host(s) The virus-infected small cardamom plants exhibit symptoms on the youngest leaf of affected tiller as slender chlorotic flecks measuring 2–5 mm in length. Later these flecks develop into pale green discontinuous stripes. These stripes run parallel to veins from midrib to leaf margins. All subsequently emerging leaves show characteristic mosaic symptoms with stripes of green tissue almost evenly distributed over the entire lamina. Often mosaic-type of mottling is seen on leaf sheaths and young leaf shoots (Uppal et al. 1945; Siddaramaiah et al. 1986). Transmission The virus is transmitted by aphid vectors Pentalonia nigronervosa, F. caladii, Aphis craccivora, A. gossypii, A. nerii, A. rumicis, Brachycaudus helichrysi, Greenidia artocarpi, Macrosiphum pisi, M. rosaeformis, M. sonchi, Schizaphis cyperi, and S. graminum in a non-persistent manner (Rao and Naidu 1974; Venugopal et al. 1997). The virus is transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants nor by seed. Use of planting material from infected plants is the primary source of spread. Virion properties and genome The virions are non-enveloped, flexuous filaments, 650 nm long, and 13–16 nm in diameter. The genome consists of a single molecule of linear, positive-sense ssRNA of 8249 nt (MF622947) (Dimitman et al. 1984; Gonsalves et al. 1986; Jacob and Usha 2001, 2002; Lopez-Moya et al. 2009; Foster 2011; Wylie et al. 2017).

References Biju CN, Siljo A, Bhat AI (2010) Survey and RT-PCR based detection of Cardamom mosaic virus affecting small cardamom in India. Indian J Virol 21:148–150 Capoor SP (1967) Katte disease of cardamom. Seminar on Cardamom. Cardamom Board, Cochin, p 25 Dimitman JE, Flores A, Nickoloff JA (1984) Cardamom mosaic – a member of the Potyvirus group in Guatemala (Abstr.). Phytopathology 74:844 Foster GD (2011) Macluravirus. Potyviridae. In: The Springer index of viruses. Springer, New York, pp 1421–1424. https://doi.org/10.1007/978-0-387-95919-1_234 Gonsalves D, Trujillo E, Hoch HC (1986) Purification and some properties of a virus associated with cardamom mosaic, a new member of Potyvirus group. Plant Dis 70:65–69 Jacob T, Usha R (2001) 30 -terminal sequence analysis of the RNA genome of the Indian isolate of cardamom mosaic virus: a new member of genus Macluravirus of Potyviridae. Virus Genes 23:81–88 Jacob T, Usha R (2002) Expression of Cardamom mosaic virus coat protein in Escherichia coli and its assembly into filamentous aggregates. Virus Res 86:133–141 Jacob T, Jebasingh T, Venugopal MN, Usha R (2003) High genetic diversity in the coat protein and 30 untranslated regions among geographical isolates of Cardamom mosaic virus from South India. J Biol Sci 25:589–595

Eleusine coracana (Finger millet)

947

Lopez-Moya JJ, Valli A, Garcia JA (2009) Potyviridae. In: Encyclopaedia of Life Sciences (ELS). NS Iacobellis, A Collmer, SW Hutcheson, JW Mansfield, CE Morris, J Murillo, NW Schaad D, E Stead, G Surico, and M Ullrich, (eds.). Kluwer Academic, Dordrecht, The Netherlands. https://doi.org/10.1002/9780470015902.a0000755.pub2 Rao DG (1977) Katte disease of small cardamom. J Plant Crop 5:23–27 Rao DG, Naidu R (1973) Studies on “Katte” or “mosaic” disease of small cardamom. J Plant Crop 1:129–136 Rao DG, Naidu R (1974) Additional vectors of katte disease of small cardamom. Indian J Hortic 31:380–381 Saigopal DVR, Naidu R, Thomas J (1990) Early detection of ‘Katte’ disease of small cardamom through ELISA. J Plant Crops 20:73–75 Siddaramaiah AL, Chandrashekar SC, Balachandra CK, Pattanshetti V (1986) Arrowroot (Marantha arundinacea L.) a new collateral host for ‘Katte’ disease of cardamom (Elettaria cardamom Maton.). Curr Sci 55:18 Siljo A, Bhat AI (2014) Reverse transcription loop-mediated isothermal amplification assay for rapid and sensitive detection of Banana bract mosaic virus in cardamom (Elettaria cardamomum). Eur J Plant Pathol 138(2):209–214 Siljo A, Bhat AI, Biju CN, Venugopal MN (2011) Occurrence of banana bract mosaic virus on cardamom. Phytoparasitica 40:77 Siljo A, Bhat AI, Biju CN, Venugopal MN (2012) Occurrence of Banana bract mosaic virus on cardamom. Phytoparasitica 40:77–85 Siljo A, Bhat AI, Biju CN (2014) Detection of Cardamom mosaic virus and Banana bract mosaic virus in cardamom using SYBR green based reverse transcription-quantitative PCR. Indian J Virol 25:137–141 Uppal BN, Varma PM, Capoor SP (1945) A mosaic disease of cardamom. Curr Sci 14:208–209 Venugopal MN (1995) Viral diseases of cardamom (Elettaria cardamommum Maton) and their management. J Spices Aromat Crops 4(1):32–39 Venugopal MN, Saju KA, Mathew MJ (1997) Primary spread of cardamom mosaic virus under different field situations. In: National Symposium on Resurgence of Vector Borne Viral Diseases, 1–3 August 1997, Gujarath Agricultural University, Anand, p 15 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. Journal of General Virology 98:352–354

Eleusine coracana (Finger millet) Family: Poaceae

Cereal

Finger millet mosaic virus

(FMMV)

Synonyms Eleusine mosaic virus Taxonomic position FMMV is a tentative member of the family Rhabdoviridae. Geographical distribution FMMV was first reported in plants of Eleusine coracana by Rao et al. (1965) from India. The virus spreads in Southern India. Symptoms and host(s) The virus-infected finger millet plants exhibit streaking, mosaic, mottling, and general chlorosis symptoms. The host range restricted to species of the Gramineae. Transmission The virus is transmitted by the delphacid planthopper vector, Sogatella longifurcifera. The virus is not transmissible by mechanical sap-inoculation.

E

948

Eleusine indica (Indian goosegrass)

Virion properties and genome The virions are rhabdo- or bullet-shaped, with a modal length of 285 nm and 80 nm wide.

Maize chlorotic mottle virus Taxonomic position Genus: Machlomovirus

(MCMV)

Family: Tombusviridae

MCMV infection in plants of Eleusine coracana was reported from Kenya (Kusia et al. 2015). This virus was also observed in Congo, China, Taiwan, and Nebraska (USA). The virus-infected finger millet plants exhibit symptoms of chlorotic mottle from the base of young leaf whorling upward to the tips and necrosis from leaf margins to the midrib. The virus is transmitted by the thirps vector, Frankliniella williamsi, and is also transmissible by mechanical sap-inoculation. For more details of MCMV, refer to Zea mays.

Sugarcane mosaic virus Taxonomic position Genus: Potyvirus

(SCMV)

Family: Potyviridae

SCMV infection in plants of Eleusine coracana was reported from Kenya (Kusia et al. 2015). The virus-infected finger millet plants exhibit symptoms of chlorotic mottle from the base of young leaf whorling upward to the tips and necrosis from leaf margins to the midrib. The virus is transmitted by the aphid vectors, Rhopalosiphum spp. and Sitobion avenae, in a non-persistent manner, by mechanical sap-inoculation, by grafting, and by seed. For more details of SCMV, refer to Saccharum officinarum.

References Kusia ES, Subramanian S, Nyasani JO, Khamis F, Villinger J, Ateka EM, Pappu HR (2015) First report of lethal necrosis disease associated with co-infection of finger millet with Maize chlorotic mottle virus and Sugarcane mosaic virus in Kenya. Plant Dis 99:899–900 Rao DG, Varma PM, Capoor SP (1965) Studies of mosaic disease of Eleusine in the Deccan. Indian Phytopath 18:139–150

Eleusine indica (Indian goosegrass) Family: Poaceae

Weed host

Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

Elymus trachycaulus (Wild rye)

949

IYSV infection in plants of Eleusine indica was reported from Zimbabwe (Karavina and Gubba 2017). The virus is transmitted by onion thrips, Thrips tabaci, in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of IYSV, refer to Iris spp. or Allium cepa.

References Karavina C, Gubba A (2017) Amaranthus sp. and Eleusine indica are natural hosts of Iris yellow spot virus in Zimbabwe. Plant Dis 101:262

E Eleusine tristachya (Spike goosegrass) Family: Poaceae

Forage crop

Wheat streak mosaic virus Taxonomic position Genus: Tritimovirus

(WSMV)

Family: Potyviridae

WSMV infection in plants of Eleusine tristachya was reported from Australia (Ellis et al. 2004). The virus-infected spike goosegrass plants exhibit leaf streaking and mottling symptoms. The virus is transmitted by the wheat curl mite (Aceria tosichella) in a semi-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of WSMV, refer to Triticum aestivum.

References Ellis MH, Rebetzke GJ, Kelman WM, Moore CS, Hyles JE (2004) Detection of wheat streak mosaic virus in four pasture grass species in Australia. Plant Pathol 53:239

Elymus trachycaulus (Wild rye) Family: Poaceae

Poa semilatent virus Taxonomic position Genus: Hordeivirus

Forage crop

(PSLV)

Family: Virgaviridae

PSLV infection in plants of Elymus trachycaulus was reported from Canada by Slykhuis (1972). The virus spreads in Canada (Alberta). The virus-infected wild rye plants show either faint leaf chlorosis or

950

Emilia sonchifolia (Lilac tasselflower)

are symptomless. The virus is transmissible by mechanical inoculation but not transmitted by seed. For more details of PSLV, refer to Poa spp.

References Slykhuis JT (1972) Poa semilatent virus from native grasses. Phytopathology 62:508–513

Emilia sonchifolia (Lilac tasselflower) Family: Asteraceae

Medicinal

Emilia yellow vein Thailand virus Taxonomic position Genus: Begomovirus

(EYVTHV)

Family: Geminiviridae

Geographical distribution EYVTHV infection in plants of Emilia sonchifolia was reported from Thailand (Zhao et al. unpublished – KY373213). Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2746 nt (KY373213 = NC_035473) (Brown et al. 2015; Zerbini et al. 2017).

Emilia yellow vein virus Taxonomic position Genus: Begomovirus

(EmYVV)

Family: Geminiviridae

Geographical distribution EmYVV infection in plants of Emilia sonchifolia was reported from China (Yang et al. 2008). Symptoms and host(s) The virus-infected lilac tasselflower plants exhibit yellow vein symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner.

Epidendrum spp.

951

Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single-coat protein. The genome consists of a single molecule of circular, single-stranded DNA (DNA-A) of 2725 nt (EU377539 = NC_010307) (Briddon 2001; Yang et al. 2008; Brown et al. 2015; Zerbini et al. 2017).

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Yang CX, Cui GJ, Zhang J, Weng XF, Xie LH, Wu ZJ (2008) Molecular characterization of a distinct begomovirus species isolated from Emilia sonchifolia. J Plant Pathol 90:475–478 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. Journal of General Virology 98:131–133

Ensete superbum (Ensete) Family: Musaceae

Edible fruit

Banana bunchy top virus Taxonomic position Genus: Babuvirus

(BBTV)

Family: Nanoviridae

BBTV infection in plants of Ensete superbum was reported from India (Selvarajan and Balasubramanian 2013). The virus-infected ensete plants exhibit symptoms of intermittent dark green dots, dash, and streaks of variable length like “Morse code” pattern on leaf sheath, midrib, leaf veins and petioles, and bunching of shorter and narrow leaves. The virus is transmitted by the banana black aphid, Pentalonia nigronervosa, in a persistent but not a propagative manner. The virus is not mechanically sap-transmissible. For more details of BBTV, refer to Musa spp.

References Selvarajan R, Balasubramanian V (2013) Natural occurrence of Banana bunchy top virus in Ensete superbum in India. Indian J Virol 24:97–98

Epidendrum spp. Synonyms Encyclia spp. Family: Orchidaceae

Ornamental

E

952

Epidendrum spp.

Cymbidium mosaic virus Taxonomic position Genus: Potexvirus

(CymMV)

Family: Alphaflexiviridae

CymMV infection in plants of Epidendrum spp. was reported from Puerto Rico, Ecuador, Florida, India, and Hawaii (Hu et al. 1993; Elliott et al. 1996; Sherpa et al. 2003; Pant et al. 2010). The virusinfected epidendrum plants exhibit symptoms of chlorotic rings with a necrotic spot in the center and bronzing of leaves. On the lower surface of the infected leaves sunken, brown or black, round to oval spots are noticed. The virus is mechanically sap-transmissible and also by contact between plants. No vector transmission is reported. For more details of CymMV, refer to Cymbidium spp.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Epidendrum spp. was reported from the USA (Hammond et al. 2011). The virus-infected epidendrum plants exhibit necrotic spot symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Odontoglossum ringspot virus Taxonomic position Genus: Tobamovirus

(ORSV)

Family: Virgaviridae

ORSV infection in plants of Epidendrum spp. was reported from India and Hawaii (Hu et al. 1993; Pant et al. 2010). The virus-infected epidendrum plants exhibit symptoms of chlorotic rings with necrotic spot in the center and bronzing of leaves. The virus is transmitted by means not involving a vector. The virus is transmissible by mechanical sap-inoculation, and also by contact between plants. For more details of ORSV, refer to Odontoglossum grande.

References Elliott MS, Zettler FW, Zimmerman MT, Barnett OW Jr, Le Grande MD (1996) Problems with interpretation of serological assays in a virus survey of orchid species from Puerto Rico, Ecuador, and Florida. Plant Dis 80:1160–1164 Hammond J, Jordan R, Vaira AM (2011) Virus diseases of ornamentals, in soils, plant growth and crop production. In: UNESCO-EOLSS Joint Committee (ed) Encyclopedia of life support systems (EOLSS), developed under the auspices of the UNESCO. Eolss Publishers, Oxford. http://www.eolss.net Hu J, Ferreira S, Wang M, Xu MQ (1993) Detection of Cymbidium mosaic virus, Odontoglossum ringspot virus, Tomato spotted wilt virus, and potyviruses infecting orchids in Hawaii. Plant Dis 77:464–468 Pant RP, Mrinal Das KB, Pun KB, Ramachandran P, Medhi RP (2010) Occurrence of Cymbidium mosaic and Odontoglossum ringspot viruses in orchid germplasm of Sikkim and Darjeeling hills, their identification and diagnosis. Indian Phytopathol 63:326–332 Sherpa AR, Hallan V, Ram R, Vij SP, Pathak P, Garg ID, Zaidi AA (2003) First report of Cymbidium mosaic virus (CymMV) in orchids from India. Plant Pathol 52:788

Epimedium spp. (Barrenwort)

953

Epilobium angustifolium (Fireweed) Synonyms Chamaenerion angustifolium Family: Onagraceae

Turnip mosaic virus Taxonomic position Genus: Potyvirus

Medicinal plant

(TuMV)

Family: Potyviridae

TuMV infection in plants of Epilobium angustifolium was reported from south-western part of Iceland (Kegler et al. 1998). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap inoculation to a large number of host plants. For more details of TuMV, refer to Brassica rapa.

References Kegler H, Ehrig F, Rabenstein F (1998) Evidence of a plant virus in Iceland. Arch Phytopath Pflanz 31:241–246

Epimedium spp. (Barrenwort) Family: Berberidaceae

Medicinal

Tobacco rattle virus

(TRV)

Taxonomic position Genus: Tobravirus

Family: Virgaviridae

TRV infection in plants of Epimedium spp. was reported from Ohio (USA) (Lockhart 2000; Fisher and Bell 2014). The virus-infected barrenwort plants exhibit symptoms of yellow ringspotting and concentric line patterns. Even mottle and mosaic symptoms are noticed in some species. Symptoms either persist, vary seasonally, or disappear soon after infection depending on the host species and virus strain. The virus is transmitted by nematode vectors of the genera Trichodorus and Paratrichodorus and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tomato mosaic virus Taxonomic position Genus: Tobamovirus

(ToMV)

Family: Virgaviridae

E

954

Epiphyllum spp. (Orchid cacti)

ToMV infection in plants of Epimedium spp. was reported from the USA (Nameth et al. 1991). The virus-infected barrenwort hybrids show mosaic, line patterns, or concentric ringspots. No vector is involved in the spread of this virus. This virus is transmissible by mechanical sap-inoculation, transmissible by grafting, and also by contact between plants. For more details of ToMV, refer to Solanum lycopersicum.

References Fisher JR, Bell J (2014) Tobacco rattle virus is detected in established landscale plantings of Epimedium sp. in Ohio. Plant Health Prog 15(1):53–54 Lockhart BEL (2000) Dicentra, Epimedium and Heuchera: new perennial ornamental hosts of Tobacco rattle virus in the United States. Plant Dis 84:1344 Nameth ST, Putnam ML, Valverde RA, Reddick BB (1991) A tobamovirus associated with mosaic and ring spot symptoms on Barrenwort (Epimedium sp.). Hortic Sci 26:1505–1506

Epiphyllum spp. (Orchid cacti) Family: Cactaceae

Orchid

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

BWYV infection in plants of Epiphyllum spp. was reported from the USA (Eicholtz et al. 2018). The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by neither mechanical sap-inoculation nor seed. For more details of BWYV, refer to Beta vulgaris.

References Eicholtz M, Grinstead S, Wu L-P, Kinarg G, Li R (2018) First report of Beet western yellows virus infecting Epiphyllum spp. Plant Dis 102:464

Episcia spp. Family: Gesneriaceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(INSV)

Eragrostis spp. (Lovegrass)

955

INSV infection in plants of Episcia capreata was reported from Italy (Roggero et al. 1999). The virusinfected episcia plants show symptoms of necrotic spots on stems and leaves. The virus is transmitted by the thrips vector, Frankliniella occidentalis, in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV was reported from a few cultivated plants of Episcia spp. in the USA (Zettler and Nagel 1983). The symptoms on the foliage of infected episcia plants were inconspicuous. There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

References Roggero P, Ciuffo M, Dellavalle G, Gotta P, Gallo S, Peters D (1999) Additional ornamental species as hosts of Impatiens necrotic spot tospovirus in Italy. Plant Dis 83:967 Zettler FW, Nagel J (1983) Infection of cultivated Gesneriads by two strains of Tobacco mosaic virus. Plant Dis 67:1123–1125

Eragrostis spp. (Lovegrass) Family: Poaceae

Forage crop

Barley yellow dwarf virus PAV Taxonomic position Genus: Luteovirus

(BYDV-PAV)

Family: Luteoviridae

BYDV-PAV infection in plants of Eragrostis curvula was reported from Western Australia (Jones et al. 1990). The virus is transmitted by a number of aphid species in a circulative, non-propagative manner. No mechanical transmission of this virus is reported. For more details of BYDV-PAV, refer to Hordeum vulgare.

Cereal yellow dwarf virus RPV Taxonomic position Genus: Polerovirus

(CYDV-RPV)

Family: Luteoviridae

CYDV-RPV infection in plants of Eragrostis curvula was reported from Western Australia (Jones et al. 1990). The virus is transmitted by a number of aphid species in a circulative, non-propagative manner.

E

956

Eragrostis spp. (Lovegrass)

No mechanical transmission of this virus is reported. For more details of CYDV-RPV, refer to Hordeum vulgare.

Eragrostis curvula streak virus Taxonomic position Genus: Eragrovirus

(ECSV)

Family: Geminiviridae

Geographical distribution ECSV infection in plants of Eragrostis curvula was reported from South Africa (Varsani et al. 2009). Symptoms and host(s) The virus-infected lovegrass plants exhibit mild streak symptoms. Transmission There is no known vector for this virus. Virion properties and genome The structure of ECSV particles has not been investigated. In common with all geminiviruses, the virions of ECSV are likely geminate (twinned icosahedra). The genome of ECSV consists of a single component of circular, single-stranded DNA of ~2754 nt (FJ665631 = NC_012664; FJ665634) (Varsani et al. 2009, 2014; Zerbini et al. 2017). The genome of ECSV is predicted to encode four gene products, two in each orientation. The expression and functions of the gene products has not been further investigated. The genome of ECSV also has an unusual nonnucleotide sequence in the predicted hairpin structure which forms part of the origin of virion-strand DNA replication: “TAAGATTCC” rather than the more usual “TAATATTAC” of other geminiviruses.

Eragrostis minor streak virus Taxonomic position Genus: Mastrevirus

(EMSV)

Family: Geminiviridae

Geographical distribution EMSV infection in plants of Eragrostis minor was reported from Africa (Martin et al. 2011). Symptoms and host(s) The virus-infected lovegrass plants display discontinuous chlorotic streaks running along the major leaf veins. Transmission The insect vector of EMSV has not been identified. In common with other viruses of the genus Mastrevirus, the vector will likely be a leafhopper. Virion properties and genome The structure of EMSV particles has not been investigated. In common with all geminiviruses, the virions of EMSV are likely geminate (twinned icosahedra).

Eragrostis spp. (Lovegrass)

957

The genome of EMSV consists of a single component of circular, single-stranded DNA of 2689 nt (JF508490 = NC_015553; KM230033, KM230032) (Palmer and Rybicki 1998; Boulton and Davies 2011; Martin et al. 2011; Muhire et al. 2013; Zerbini et al. 2017). In common with all mastreviruses, EMSV is predicted to encode four gene products, two in each orientation. Based on sequence homology to previously studied mastreviruses, these would code for the coat protein and movement protein in the virion-sense and in the complementary-sense for the replication-associated protein (Rep) and Rep A.

Eragrostis streak virus

(ESV)

E

Taxonomic position Genus: Mastrevirus

Family: Geminiviridae

Geographical distribution ESV infection in plants of Eragrostis was reported from Zimbabwe (Shepherd et al. 2008). Symptoms and host(s) The virus-infected lovegrass plants exhibit streaking symptoms on the leaves. Transmission The virus is transmitted by leafhopper vectors in a persistent, circulative, and non-propagative manner. The virus is not mechanically sap-transmissible. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm, with a single coat protein. The genome is a monopartite, circular, and single-stranded DNA of 2746 nt (EU244915 = NC_010352) (Palmer and Rybicki 1998; Shepherd et al. 2008; Boulton and Davies 2011; Muhire et al. 2013; Zerbini et al. 2017).

Sugarcane mosaic virus Taxonomic position Genus: Potyvirus

(SCMV)

Family: Potyviridae

SCMV infection in plants of Eragrostis exasperata was reported from Kenya (Louie 1980). The virusinfected lovegrass plants exhibit chlorotic streaking symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of SCMV, refer to Saccharum officinarum.

Wheat streak mosaic virus Taxonomic position Genus: Tritimovirus

(WSMV)

Family: Potyviridae

WSMV infection in plants of Eragrostis curvula was reported from Australia (Ellis et al. 2004). The virus-infected lovegrass plants exhibit leaf streaking and mottling symptoms. The virus is transmitted

958

Erechtites spp. (Burnweeds)

by the wheat curl mite (Aceria tosichella) in a semi-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of WSMV, refer to Triticum aestivum.

References Boulton MI, Davies JW (2011) Mastrevirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 597–603. https://doi.org/10.1007/978-0-387-95919-1_83 Ellis MH, Rebetzke GJ, Kelman WM, Moore CS, Hyles JE (2004) Detection of wheat streak mosaic virus in four pasture grass species in Australia. Plant Pathol 53:239 Jones RAC, McKirdy SJ, Shivas RG (1990) Occurrence of barley yellow dwarf viruses in over-summering grasses and cereal crops in Western Australia. Australas Plant Pathol 19:90–96 Louie R (1980) Sugarcane mosaic virus in Kenya. Plant Dis 64:944–947 Martin DP, Linderme D, Lefeuvre P, Shepherd DN, Varsani A (2011) Eragrostis minor streak virus: an Asian streak virus in Africa. Arch Virol 156:1299–1303 Muhire B, Martin DP, Brown JK, Navas-Castillo J, Moriones E, Zerbini FM, Rivera-Bustamante R, Malathi VG, Briddon RW, Varsani A (2013) A genome-wide pairwise-identity-based proposal for the classification of viruses in the genus Mastrevirus (family Geminiviridae). Arch Virol 158(6):1411–1424 Palmer KE, Rybicki EP (1998) The molecular biology of mastreviruses. Adv Virus Res 50:183–234 Shepherd DN, Varsani A, Windram OP, Lefeuvre P, Monjane AL, Owor BE, Martin DP (2008) Novel sugarcane streak and sugarcane streak reunion mastreviruses from southern Africa and La Reunion. Arch Virol 153:605–609 Varsani A, Shepherd DN, Dent K, Monjane AL, Rybicki EP, Martin DP (2009) A highly divergent South African geminivirus species illuminates the ancient evolutionary history of this family. Virol J 6:36 Varsani A, Navas-Castillo J, Moriones E, Hernandez-Zepeda C, Idris A, Brown JK, Zerbini FM, Martin DP (2014) Establishment of three new genera in the family Geminiviridae: Becurtovirus, Eragrovirus and Turncurtovirus. Arch Virol 159:2193–2203 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. Journal of General Virology 98:131–133

Erechtites spp. (Burnweeds) Family: Asteraceae

Bidens mottle virus Taxonomic position Genus: Potyvirus

Weed host

(BiMoV)

Family: Potyviridae

BiMoV infection in plants of Erechtites hieraciifolius was reported from the USA (Purcifull and Zitter 1973). The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BiMoV, refer to Bidens spp.

Erectites yellow mosaic virus Taxonomic position Genus: Begomovirus

(ErYMV)

Family: Geminiviridae

Eremurus spp.

959

Geographical distribution ErYMV infection in plants of Erechtites valerianifolia was reported from Vietnam (Ha et al. 2008). Symptoms and host(s) The virus-infected burnweed plants exhibit variegation and yellow vein symptoms. Transmission The transmission of ErYMV has not been investigated. It is likely that, in common with other begomoviruses, ErYMV will be transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Virion properties and genome The structure of ErYMV particles has not been investigated. In common with all geminiviruses, the virions of ErYMV are likely geminate (twinned icosahedra). ErYMV is a typical Old World monopartite begomovirus. The genome of ErYMV consists of a single circular molecule of a single-stranded DNA of 2751 nt (DQ641698 = NC_009549) (Briddon 2001; Brown et al. 2015; Zerbini et. al 2017). The characterized genome of ErYMV encodes the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated for OYCrV. ErYMV associates with a betasatellite of 1342 nt (DQ641713) (Ha et al. 2008).

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Ha C, Coombs S, Revill P, Harding R, Vu M, Dale J (2008) Molecular characterization of begomoviruses and DNA satellites from Vietnam: additional evidence that the New World geminiviruses were present in the Old World prior to continental separation. J Gen Virol 89:312–326 Purcifull DE, Zitter TA (1973) A serological test for distinguishing bidens mottle and lettuce mosaic viruses, Florida Agricultural Experiment Stations Journal Series No. 5132, pp 143–145 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. Journal of General Virology 98:131–133

Eremurus spp. Family: Xanthorrhoeaceae

Tobacco rattle virus Taxonomic position Genus: Tobravirus

Ornamental

(TRV)

Family: Virgaviridae

E

960

Erica spp.

TRV infection in plants of Eremurus spp. was reported from Hungary (Nemethy 1994). The virusinfected eremurus plants exhibit symptoms of yellow green or gray necrotic spots on the leaves. The virus is transmitted by nematode vectors and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

References Nemethy IZ (1994) Survey on virus diseases of bulbous flowers in Hungary. Acta Hortic 377:267–274

Erica spp. Family: Ericaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Erica spp. was reported from Iran (Ghotbi and Shahraeen 2012). The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV was reported in plants of Erica spp. in Iran (Ghotbi and Shahraeen 2012; Ghotbi 2013). The virus-infected Erica plants exhibit symptoms of necrotic leaf spot, chlorosis, and stunting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Erica spp. was reported from Iran (Ghotbi and Shahraeen 2012). The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Eruca sativa (Arugula)

961

References Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iran J Plant Pathol 49:41 Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses affecting propagated ornamentals in Northern Iran. Int Res J Microbiol 3:373–381

Erodium moschatum (Whitestem filaree) Family: Geraniaceae

Weed host

Turnip mosaic virus Taxonomic position Genus: Potyvirus

E

(TuMV)

Family: Potyviridae

TuMV infection in plants of Erodium moschatum was reported from New Zealand (Ochoa Corona et al. 2007). The virus-infected whitestem filaree plants exhibit chlorosis, mild to severe mosaic, and necrosis symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

References Ochoa Corona FM, Lebas BSM, Elliott DR, Tang JZ, Alexander BJR (2007) New host records and new host family range for Turnip mosaic virus in New Zealand. Aust Plant Dis Notes 2:127–130

Eruca sativa (Arugula) Family: Brassicaceae

Leafy vegetable

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Eruca sativa was reported from Austria (Grausgruber-Groger 2012). The virus-infected arugula plants exhibit necrotic spot symptoms on the leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

962

Radish mosaic virus Taxonomic position Genus: Comovirus

Eruca sativa (Arugula)

(RaMV)

Family: Secoviridae

RaMV infection in plants of Eruca sativa was reported from Saudi Arabia and Egypt (Milne et al. 1980; Zein and Shafie 2005). The virus-infected arugula plants exhibit mosaic, ringspots, leaf distortion, veinal necrosis, and systemic necrosis symptoms. The virus is transmitted by beetle vectors and is also transmissible by mechanical sap-inoculation, and grafting. For more details of RaMV, refer to Raphanus sativus.

Tomato chlorosis virus Taxonomic position Genus: Crinivirus

(ToCV)

Family: Closteroviridae

ToCV infection in plants of Eruca sativa was reported from South America (Boiteux et al. 2016). The virus-infected arugula plants exhibit symptoms of interveinal yellowing in the lower leaves. The virus is transmitted by whitefly vectors in a semi-persistent manner. For more details of ToCV, refer to Solanum lycopersicum.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Eruca sativa was reported from Saudi Arabia and Brazil (Al-Saleh et al. 2009; Ribeiro-Junior et al. 2018). The virus-infected arugula plants exhibit mosaic, stunting, and deformation symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

Turnip ringspot virus

(TuRSV)

Taxonomic position TuRSV is a tentative member of the genus Comovirus and family Secoviridae. TuRSV infection in plants of Eruca sativa was reported from Brazil (Picoli et al. 2012). The virusinfected arugula plants exhibit leaf deformation and mosaic symptoms. The virus is mechanically saptransmissible. For more details of TuRSV, refer to Brassica rapa.

Eryngium spp.

963

References Al-Saleh MA, Al-Shahwan IM, Abdalla OA, Amer MA (2009) Identification and coat protein nucleotide sequence of Turnip Mosaic Potyvirus from Eruca sativa in Saudi Arabia. Asian J Plant Pathol 3:27–38 Boiteux LS, Fonseca MEN, Reis A, Costa AF, Fontes MG, Gonzalez-Arcos M (2016) Wild radish (Raphanus species) and garden rocket (Eruca sativa) as new Brassicaceae Hosts of Tomato chlorosis virus in South America. Plant Dis 100:1027 Grausgruber-Groger S (2012) First report of Impatiens necrotic spot virus on Ocimum basilicum, Eruca sativa and Anthriscus cerefolium in Austria. New Dis Rep 26:12 Milne RG, Masenga V, Lenzi R, Lovisolo O (1980) Radish mosaic virus in Eruca sativa miller. Phytopathol Mediterr 19(2/3):145–149 Picoli MHS, De Souto ER, Da Silva JM, Carnelossi PR, Garcia A, Almedia AMR (2012) A Comovirus infecting rocket (Eruca sativa) in Brasil. J Phytopathol 160:55–57 Ribeiro-Junior MR, Baldini LFS, Nozaki DN, Cruciol GCD, Pantoja KFC, Marchi BR, Moura MF, Pavan MA, KrauseSakate R (2018) Biological and molecular characterization of a basal-Brassica/Raphanus Turnip mosaic virus isolate from Eruca sativa. Trop Plant Pathol. https://doi.org/10.1007/s40858-017-0207-8 Zein SN, Shafie MS (2005) Radish Mosaic Comovirus (RaMV) isolated from Eruca sativa L. Egypt J Virol 2:61–76

Eryngium spp. Family: Apiaceae

Ornamental

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

ArMV infection in plants of Eryngium spp. was reported from Lithuania (Samuitiene et al. 2008). The virus-infected eryngium plants exhibit symptoms of stunting, leaves slightly distorted, and light green spots. The virus is transmitted by the nematode vectors, Xiphinema diversicaudatum and X. coxi, in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Broad bean wilt virus 1 Taxonomic position Genus: Fabavirus

(BBWV-1)

Family: Secoviridae

BBWV-1 infection in plants of Eryngium spp. was reported from Hungary (Wolf 1972). The virusinfected eryngium plants show severe mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BBWV-1, refer to Vicia faba.

E

964

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Eryngium spp.

(CMV)

Family: Bromoviridae

CMV infection in plants of Eryngium amethystinum and E. yuccifolium was reported from the Ohio (USA) (Wolf 1972; Fisher et al. 1997; Whitten and Nameth 2004). The virus-infected eryngium plants exhibit symptoms of severe mosaic, yellowing, and stunting. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Tobacco mild green mosaic virus Taxonomic position Genus: Tobamovirus

(TMGMV)

Family: Virgaviridae

TMGMV infection in plants of Eryngium spp. was reported from the USA (Johnson and Heim 1948). The virus-infected eryngium plants exhibit symptoms of foliar yellow mosaic. No insect vector is reported for this virus. The virus is transmissible by mechanical sap-inoculation, and has a wide host range. The virus is transmissible by grafting and also by contact between plants. For more details of TMGMV, refer to Nicotiana tabacum.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV was reported to infecting plants of Eryngium spp. in Hungary and Lithuania (Wolf 1972; Navalinskiene and Samuitiene 2006). The virus-infected eryngium plants exhibit severe mosaic symptoms. The virus is transmitted by nematode vectors and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

Plants of Eryngium spp. were found to be infected with ToRSV in Lithuania (Navalinskiene and Samuitiene 2000, 2006). The virus-infected eryngium plants exhibit symptoms of stunting, leaf and flower distortion, chlorotic and necrotic spots and streaks, ringspots, and vein necrosis. The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation, and by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Eucharis spp. (Eucharis candida; E. grandiflora) (Amazon lily)

965

References Fisher JR, Sanchez-Cuevas M-C, Nameth ST, Woods VL, Ellett CW (1997) First report of Cucumber mosaic virus in Eryngium amethystinum, Canna spp., and Aquilegia hybrids in Ohio. Plant Dis 81:1331 Johnson J, Hein EM (1948) A yellow-mosaic virus associated with Eryngium mosaic. Phytopathology 38:15 Navalinskiene M, Samuitiene M (2000) Natural occurrence of tomato ringspot nepovirus in ornamental plants in Lithuania. Proceedings of the International Conference: Development of environmentally friendly plant protection in the Baltic Region, Tartu, Estonia, September 28–29, pp 140–143 Navalinskiene M, Samuitiene M (2006) Dekoratyvinių augalų virusin_es ligos ir jų suk_el_ejai Lietuvoje. Lutut_e, Kaunas, 254 p Samuitien_e M, Navalinskien_e M, Jackevičien_e E (2008) Arabis mosaic virus on ornamental plants. Biologija 54:264–268 Whitten KR, Nameth SGP (2004) First Report of Cucumber mosaic virus in Eryngium yuccifolium (Rattlesnake Master) in Ohio. Plant Dis. 88:1384 Wolf P (1972) Virus diseases of the genus Eryngium. Acta Phytopathol Acad Sci Hung 7:363–367

Erythrina spp. Family: Fabaceae

Trees/Shrubs

Pepper golden mosaic virus Taxonomic position Genus: Begomovirus

(PepGMV)

Family: Geminiviridae

PepGMV infection in plants of Erythrina spp. was reported from Costa Rica (Castro et al. 2013). The virus-infected erythrina plants exhibit yellow veins and vein-thickening symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. For more details of PepGMV, refer to Capsicum annuum.

References Castro RM, Moreira L, Rojas MR, Gilbertson RL, Hernandez E, Mora F, Ramirez P (2013) Occurrence of squash yellow mild mottle virus and Pepper golden mosaic virus in potential new hosts in Costa Rica. Plant Pathol J 29:285–293

Eucharis spp. (Eucharis candida; E. grandiflora) (Amazon lily) Family: Amaryllidaceae

Ornamental

Amazon lily mild mottle virus Taxonomic position Genus: Anulavirus

(ALiMMV)

Family: Bromoviridae

E

966

Eucharis spp. (Eucharis candida; E. grandiflora) (Amazon lily)

Geographical distribution ALiMMV infection in plants of Eucharis grandiflora was reported from Japan (Fuji et al. 2013). Symptoms and host(s) The virus-infected Amazon lily plants show mild mosaic or mottle symptoms. Transmission Thrips are involved in transmitting this virus by carrying the virus-infected pollen grains on their bodies and introducing the virus into the susceptible hosts while feeding. The virus is transmitted efficiently through seeds of Chenopodium quinoa, C. amaranticolor, and Tetragonia expansa, but not by Myzus persicae which allowed short acquisition feeds on T. expansa. Virion properties and genome The virions are non-enveloped and quasi-spherical, with a diameter ranging from 25 to 35 nm, and have a poorly resolved surface structure. The genome is segmented, with tripartite, linear, positive-sense ssRNAs composed of RNA1 (3169 nt, AB724113), RNA2 (2507 nt, AB724114), and RNA3 (2530 nt, AB724115). Each genomic segment has a 50 cap (Fuji et al. 2013).

Amazon lily mosaic virus Taxonomic position Genus: Potyvirus

(ALiMV)

Family: Potyviridae

Geographical distribution ALiMV was first reported in plants of Eucharis grandiflora by Terami et al. (1993) from Japan. This virus spreads in the Eastern Asian region, Japan, and Taiwan (Terami et al. 1995; Hu and Chang 2004). Symptoms and host(s) The virus-infected Amazon lily plants show leaf mosaic symptoms. Transmission The virus is transmitted by the aphid vector Myzus persicae in a non-persistent manner, and the virus is lost by the vector when it molts. This virus is also transmissible by mechanical sap-inoculation to the members belonging to less than three families, and susceptible plants mostly show mosaic symptoms. Virion properties and genome The virions are non-enveloped, flexuous filaments, 760 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of c.9.5–10 kb (Revers and Garcia 2015). A partial genome sequence of 1298 nt is available (AB158523) (Fuji et al. 2004; Wylie et al. 2017).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

Eucharis spp. (Eucharis candida; E. grandiflora) (Amazon lily)

967

A CMV subgroup I isolate was detected infecting plants of Eucharis gradiflora in Japan (Terami et al. 2004). The virus-infected Amazon lily plants exhibit mosaic symptoms on leaves and a slight distortion of flower petals. The virus is transmitted by a number of aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of host plants. For more details of CMV, refer to Cucumis sativus.

Hippeastrum mosaic virus Taxonomic position Genus: Potyvirus

(HiMV)

E Family: Potyviridae

HiMV infection in plants of Eucharis spp. was reported from the Netherlands, India, Taiwan, and Brazil (Jayasinghe and Dijkstra 1979; Hu and Chang 2004; Alexandre et al. 2011). The virus-infected Amazon lily leaves show severe rugose mosaic or light green striping, with some leaf deformation, and petals show light green striping symptoms. The virus is transmitted by aphid vectors in a nonpersistent manner, and is also transmissible by mechanical sap-inoculation. For more details of HiMV, refer to Hippeastrum spp.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Eucharis grandiflora was reported from Japan (Uga and Tsuda 2005). The virus-infected Amazon lily plants show typical tospoviral symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tobacco ringspot virus

(TRSV)

Synonyms Eucharis mottle virus

Taxonomic position Genus: Nepovirus

Family: Secoviridae

TRSV was first reported in plants of Eucharis candida from the USA in plants imported from Peru (Kahn and Scott 1962; Kahn et al. 1962). The virus-infected Amazon lily plants show leaf mottling symptoms. The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

968

Euonymus spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Eucharis grandiflora was reported from Sao Paulo State, Brazil, and the USA (Hausbeck et al. 1992; Alexandre et al. 2014). The virus-infected Amazon lily plants exhibit symptoms of chlorotic rings and spots on the leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Alexandre MAV, Duarte LML, Rivas EB, Cilli A, Harakava R, Galleti SR, Kitajima EW (2011) Hippeastrum mosaic virus diagnosed in Hippeastrum and Eucharis in Brazil. J Plant Pathol 93:643–649 Alexandre MAV, Duarte LML, Harakava R, Chaves ALR, Marque JM, Kitajima EW (2014) Amazon lily: a new natural host of Tomato spotted wilt virus. New Dis Rep 30:13 Fuji S, Terami F, Furuya H, Naito H, Fukumoto F (2004) Nucleotide sequence of the coat protein genes of alstroemeria mosaic virus and amazon lily mosaic virus, a tentative species of genus potyvirus. Arch Virol 149:1843–1849 Fuji S, Kikuchi M, Ueda S, Toda T, Furuya H, Fukumoto F, Hanada K (2013) Characterization of a new Anulavirus isolated from Amazon lily plants. Arch Virol 158:201–206 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among greenhouse ornamentals in Pennsylvania. Plant Dis 76:795–800 Hu WC, Chang YC (2004) A new mosaic disease of Amazon lily in Taiwan. Plant Pathol 53:240 Jayasinghe U, Dijkstra J (1979) Hippeastrum mosaic virus and another filamentous virus in Eucharis grandiflora. Neth J Plant Pathol 85:47–65 Kahn RP, Scott HA (1962) New strain of tobacco ringspot virus isolated from an unidentified imported species of Eucharis. Phytopathology 52:16 Kahn RF, Scott HA, Monroe RL (1962) Eucharis mottle strain of tobacco ringspot virus. Phytopathology 52:1211–1216 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Terami F, Honda Y, Fukumoto F (eds) (1993) A new potyvirus isolated from Eucharis grandiflora, Amazon lily mosaic virus. Ann Phytopathol Soc Jpn 59:334 Terami F, Honda Y, Fukumoto Y (1995) Amazon lily mosaic virus, a new potyvirus infecting Amazon lily (Eucharis grandiflora). Ann Phytopathol Soc Jpn 61:1–6 Terami F, Fukumoto F, Hanada K (2004) Cucumber mosaic virus isolated from Amazon lily (Eucharis grandiflora). J Gen Plant Pathol 70:192–193 Uga H, Tsuda S (2005) A one-step reverse transcription-polymerase chain reaction system for the simultaneous detection and identification of multiple tospovirus infections. Phytopathology 95:166–171 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. Journal of General Virology 98:352–354

Euonymus spp. Family: Celastraceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Ornamental

(AMV)

Family: Bromoviridae

Euonymus spp.

969

AMV was reported in plants of Euonymus japonica var. microphylla from Italy (Bellardi et al., 1994; Turina et al. 1994). Younger leaves of infected euonymus plants had chlorotic ring and line patterns, while older leaves had chlorotic blotches and concentric necrotic rings. The virus is transmitted by a large number of aphid species in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of AMV, refer to Medicago sativa.

Cherry leaf roll virus Taxonomic position Genus: Nepovirus

(CLRV)

Family: Secoviridae

CLRV infection was reported affecting plants of E. europaeus in the USA (Larsen et al. 1990). The virus-infected euonymus plants show leaf mottling, interveinal chlorosis, stunting, dieback, and witches’ broom symptoms. The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation, and by grafting. For more details of CLRV, refer to Prunus avium.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV was reported from plants of Euonymus japonica var. microphylla in the USA (Barnett and Baxter 1974). The virus-infected euonymus plants exhibit symptoms of white mosaic in the leaves. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Euonymus fasciation virus

(EFV)

Taxonomic position EFV is a tentative member of the family Rhabdoviridae. Geographical distribution EFV was first reported in plants of Euonymus japonica from France by Codaccioni and Cossard (1977). Symptoms and host(s) The virus-infected euonymus plants exhibit fascinated stems, foliar spotting, and vein yellowing symptoms. Virion properties and genome The virions are enveloped, bacilliform virions, c.70  300 nm (Plese and Eric 1980) or 45–100 nm in diameter and 130–300 nm long (King et al. 2012).

E

970

Plum pox virus Taxonomic position Genus: Potyvirus

Euonymus spp.

(PPV)

Family: Potyviridae

PPV infection in plants of European spindle tree (Euonymus europaeus) was reported from the Czech Republic (Polak 2001). The virus-infected euonymus plants exhibit symptoms of distinct diffuse foliar spots and rings. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of PPV, refer to Prunus domestica.

Strawberry latent ringspot virus Taxonomic position Genus: Unassigned

(SLRSV)

Family: Secoviridae

SLRSV infection in plants of Euonymus europaeus was reported from Germany (Schmelzer 1969). The virus-infected euonymus plants show yellow mottle of the leaves. The virus is transmitted by the nematode vectors (Xiphinema spp.) and is also transmissible by mechanical sap-inoculation, and by grafting. For more details of SLRSV, refer to Fragaria spp.

Tobacco necrosis virus D

(TNV-D)

Synonyms Euonymus mosaic virus Taxonomic position Genus: Betanecrovirus

Family: Tombusviridae

TNV-D infection in plants of Euonymus europaeus was reported from Czechoslovakia and Slovakia (Bojnansky and Kosljarova 1968; Subikova 1973; Subikova et al. 1974; Mali 1976). The virus-infected euonymus plants show foliar mosaic, greenish-yellow, or yellowish irregular spots. “Euonymus mosaic virus” was shown to be a strain of TNV-D, on the basis of serology and infectivity to diagnostic hosts. The virus is transmitted by a fungal vector, Olpidium brassicae (Mali 1976). For more details of TNVD, refer to Nicotiana tabacum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Euonymus fortunei var. vegetus and E. kiautschovicus was reported from the USA (Puffinberger and Corbett 1985). The virus-infected euonymus plants display chlorotic rings and oakleaf pattern symptoms. The virus is transmitted by a nematode vector Xiphinema americanum in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ToRSV, refer to Solanum lycopersicum.

Eupatorium spp.

971

References Barnett OW, Baxter LW (1974) Cucumber mosaic virus in Nandina and other ornamentals. Acta Hortic 36:65–74 Bellardi MG, Turina M, Rubies-Autonell C (1994) Virus diseases of ornamental shrubs. VIII. Euonymus japonica var. microphylla: a new host of alfalfa mosaic virus (AMV). Phytopathol Mediterr 33:230–233 Bojnansky V, Kosljarova V (1968) Euonymus mosaic. Biol Plant 10(4):322–324 Codaccioni M, Cossard C (1977) Transmission experimentale de particles du type rhabdovirus, naturellement presentes dans les tiges fasciees dune variete de Fusain. Cr Hebd Seanc Acad Sci Paris 284:701 King AMQ, Adams MJ, Carstens EB, Lefkowitz EJ (2012) Virus taxonomy: classification and nomenclature of viruses, ninth report of the international committee on taxonomy of viruses. Elsevier, San Diego Larsen RC, Gergerich RC, Kim KS (1990) Characterization and ultrastructural studies of a nepovirus from Euonymus. Phytopathology 80:134–140 Mali VR (1976) Studies on Euonymus mosaic virus disease and its transmission by Olpidium brassicae in Czechoslovakia. Indian Phytopathol 29:262–268 Plese N, Eric Z (1980) Rhabdovirus in Euonymus japonica. Acta Phytopathol Acad Sci Hung 15:291–295 Polak J (2001) European spindle tree and common privet as new natural hosts of plum pox virus. Acta Hortic 550:125–128 Puffinberger CW, Corbett MK (1985) Euonymus chlorotic ringspot disease caused by tomato ringspot viris. Phytopathology 75:423–428 Schmelzer K (1969) Strawberry latent ringspot virus in Euonymus, Acacia and Aesculus. Phytopathol Z 66:1–24 Subikova V (1973) The mechanical transmission of euonymus mosaic virus, maple leaf perforation by leaf extract or leaf nucleic acid to herbaceous plants. Biol Plant 15:166–170 Subikova V, Baumgartnerova H, Bojnansky V (1974) Purification, electron microscopy and serology of virus isolated from Euonymus europaeus. Biol Plant 16(5):321–324 Turina M, Bellardi MG, Rubies-Autonell C (1994) First report of natural infection of Euonymus japonica var. microphylla by alfalfa mosaic virus. Plant Dis 78:925

Eupatorium spp. Family: Asteraceae

Ornamental

Eupatorium yellow vein mosaic virus Taxonomic position Genus: Begomovirus

(EpYVMV)

Family: Geminiviridae

Geographical distribution EpYVMV infection in plants of Eupatorium makinoi was reported from Japan (Saunders et al. 2003). Symptoms and host(s) The virus-infected eupatorium plants exhibit variegation and yellow vein symptoms. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminated) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2778 nt (AJ438937) (Saunders et al. 2003; Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). Eupatorium yellow vein mosaic betasatellite DNA molecule is associated with EpYVMV, and consists of 1359 nt (AB300464) (Ueda et al. 2008; Zhou 2013).

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Eupatorium spp.

Eupatorium yellow vein virus Taxonomic position Genus: Begomovirus

(EpYVV)

Family: Geminiviridae

Geographical distribution EpYVV infection in plants of Eupatorium makinoi was reported from Japan (Onuki and Hanada 2000; Saunders et al. 2003). This virus was previously described as a variant of Tobacco leaf curl virus (Kitamura et al. 2004).

Symptoms and host(s) The virus-infected eupatorium plants exhibit variegation and bright yellow vein symptoms. The virus has only been identified in Eupatorium makinoi and tomato (Solanum lycopersicum).

Transmission The transmission of the EpYVV has not been investigated. It is likely that, in common with other begomoviruses, EpYVV is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. Experimentally the virus has also been shown to infect Nicotiana benthamiana by Agrobacterium-mediated inoculation of the cloned virus genome in the presence of a betasatellite (Saunders et al. 2008).

Virion properties and genome The structure of the virions of EpYVV has not been investigated. In common with all geminiviruses, the virions of EpYVV are likely geminate (twinned quasi-icosahedra). EpYVV is a typical Old World monopartite begomovirus. The genome consists of a single circular molecule of single-stranded DNA of ~2766 nt (AB007990 = NC_003556; AJ438937; AB079766 AB300463, AB433979, AJ438936) (Onuki and Hanada 2000; Briddon 2001; Saunders et al. 2003; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes of EpYVV encode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated. Eupatorium yellow vein betasatellite DNA molecule is associated with EpYVV, and consists of 1356 nt (AJ438938) (Saunders et al. 2003; Zhou 2013).

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Eupatorium capillifolium and E. purpureum (Joe-Pye weed) was reported from the USA (Sabanadzovic et al. 2010). The virus-infected eupatorium plants show systemic chlorosis, mottle, and downward leaf rolling symptoms. The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Euphorbia caput-medusae (Medusa’s head)

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

973

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Eupatorium capillifolium has been reported in the USA and Greece (Groves et al. 2002; Mullis et al. 2009). The virus-infected eupatorium plants exhibit symptoms of chlorotic and necrotic rings and line patterns. The virus is transmitted by thrips vectors in a persistentpropagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Groves RL, Walgenbach JF, Moyer JW, Kennedy GG (2002) The role of weed hosts and tobacco thrips, Frankliniella fusca, in the epidemiology of Tomato spotted wilt virus. Plant Dis 86:573–582 Kitamura K, Murayama A, Ikegami M (2004) Evidence for recombination among isolates of Tobacco leaf curl Japan virus and Honeysuckle yellow vein mosaic virus. Arch Virol 149:1221–1229 Mullis S, Bertrand PF, Brown SL, Csinos A, Diaz-Perez JC, Gitaitis R, Hickman LL, Johnson A, LaHue SS, Martinez N, McPherson RM, Nischwitz C, Reay-Jones F, Riley D, Sherwood J, Stevenson K, Wells L (2009) Tospoviruses in Solanaceae and other crops in the coastal plain of Georgia. University of Georgia, College of Agricultural and Environmental Sciences, Bulletin 1354, pp 51 Onuki M, Hanada K (2000) Genomic structure of a geminivirus in the genus Begomovirus from yellow vein-affected Eupatorium makinoi. J Gen Plant Pathol 66:176–181 Sabanadzovic S, Ingram DM, Lawrence AM (2010) First report of Tobacco ringspot virus in Joe-Pye Weed (Eupatorium purpureum) in Mississippi. Plant Dis 94:126 Saunders K, Bedford ID, Yahara T, Stanley J (2003) Aetiology: the earliest recorded plant virus disease. Nature 422(6934):831 Saunders K, Briddon RW, Stanley J (2008) Replication promiscuity of DNA-b satellites associated with monopartite begomoviruses; deletion mutagenesis of the Ageratum yellow vein virus DNA-b satellite localises sequences involved in replication. J Gen Virol 89:3165–3172 Ueda S, Onuki M, Hanada K, Takanami Y (2008) Unique grouping of the Far East Asian begomovirus complex based on sequence analyses of the DNA-A genome and associated DNAbeta satellite molecules isolated from tomato, honeysuckle and Eupatorium plants in Japan. Arch Virol 153(3):417–426 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133 Zhou X (2013) Advances in understanding begomovirus satellites. Ann Rev Phytopathol 51:357–381

Euphorbia caput-medusae (Medusa’s head) Family: Euphorbiaceae

Ornamental

Euphorbia caput-medusae latent virus Taxonomic position Genus: Capulavirus

Family: Geminiviridae

(EcmLV)

E

974

Euphorbia eritrea

Geographical distribution EcmLV infection in plants of Euphorbia caput-medusae was reported from South Africa (Bernardo et al. 2013, 2016). Symptoms and host(s) The virus-infected medusa’s head plants do not exhibit any external symptoms. Transmission There is no known vector for this virus. The virus is mechanically sap-transmissible to Solanum lycopersicum and Nicotiana benthamiana (Bernardo et al. 2013). Virion properties and genome The virions are twinned (geminated) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is a monopartite circular, single-stranded DNA of 2678 nt (HF921459) (Bernardo et al. 2013; Varsani et al. 2017; Zerbini et al. 2017).

References Bernardo P, Golden M, Akram M, Naimuddin NN, Fernandez E, Granier M, Rebelo AG, Peterschmitt M, Martin DP, Roumagnac P (2013) Identification and characterisation of a highly divergent geminivirus: evolutionary and taxonomic implications. Virus Res 177(1):35–45 Bernardo P, Muhire B, François S, Deshoux M, Hartnady P, Farkas K, Kraberger S, Filloux D, Fernandez E, Galzi S, Ferdinand R, Granier M, Marais A, Monge Blasco P, Candresse T, Escriu F, Varsani A, Harkins GW, Martin DP, Roumagnac P (2016) Molecular characterization and prevalence of two capulaviruses: Alfalfa leaf curl virus from France and Euphorbia caput-medusae latent virus from South Africa. Virology 493:142–153 Varsani A, Roumagnac P, Fuchs M, Navas-Castillo J, Moriones E, Idris A, Briddon RW, Rivera-Bustamante R, Murilo Zerbini F, Martin DP (2017) Capulavirus and Grablovirus: two new genera in the family Geminiviridae. Arch Virol 162(6):1819–1831 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

Euphorbia eritrea Family: Euphorbiaceae

Ornamental

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV was reported from Italy in plants of Euphorbia eritrea (Salomone et al. 2003). The virusinfected Euphorbia eritrea plants show chlorotic and necrotic spots in the stem, starting from the attachment point of true leaves. The virus is transmitted by thrips vectors and is also transmissible by mechanical sap-inoculation to a large number of hosts (Salomone et al. 2003). For more details of TSWV, refer to Solanum lycopersicum.

Euphorbia heterophylla (Painted Euphorbia)

975

References Salomone A, Masenga V, Minuto G, Parodi C, Roggero P (2003) First report of Tomato spotted wilt virus (Tospovirus, Bunyaviridae) infecting Euphorbia eritrea and Asclepias curassavica in Liguria, Italy. Plant Pathol 52:806

Euphorbia geniculata Family: Euphorbiaceae

Weed host

Onion yellow dwarf virus Taxonomic position Genus: Potyvirus

E

(OYDV)

Family: Potyviridae

OYDV infection in plants of Euphorbia geniculata was reported from Egypt (Abdel Wahab et al. 2009). The virus-infected Euphorbia geniculata plants show yellow lesions on the leaves. The virus is transmitted by a number of aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of OYDV, refer to Allium cepa.

References Abdel Wahab AS, Elnagar S, El-Sheikh MAK (2009) Incidence of aphid-borne Onion yellow dwarf virus (OYDV) in alliaceae crops and associated weeds in Egypt. 4th conference on recent technologies in agriculture. http://conf2009. agr.cu.edu.eg/volum1/4.pdf

Euphorbia heterophylla (Painted Euphorbia) Synonyms Euphorbia prunifolia Family: Euphorbiaceae

Ornamental

Euphorbia mosaic Peru virus Taxonomic position Genus: Begomovirus

(EuMPV)

Family: Geminiviridae

Geographical distribution EuMPV infection in plants of Euphorbia heterophylla was reported from Peru (Shepherd et al. 2008).

976

Euphorbia heterophylla (Painted Euphorbia)

Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm, with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2600 nt (AM886131 = NC_039082) (Briddon 2001; Shepherd et al. 2008; Brown et al. 2015; Zerbini et al. 2017).

Euphorbia mosaic virus Taxonomic position Genus: Begomovirus

(EuMV)

Family: Geminiviridae

Geographical distribution EuMV was first reported in plants of Euphorbia heterophylla from Brazil by Costa and Bennett (1950). The virus spreads in Brazil, Puerto Rico, Peru, Mexico, Jamaica, Cuba, the USA (Florida), and Venezuela (Debrot and Centeno 1985; Hernandez-Zepeda et al. 2007; CABI/EPPO 2009; Collins et al. 2009; Gregorio-Jorge et al. 2010; Fiallo-Olive et al. 2012). Symptoms and host(s) The virus-infected painted euphorbia plants exhibit golden mosaic and leaf malformation symptoms. The virus has been isolated from Euphorbia heterophylla, Wissadula amplissima, pepper (Capsicum annuum), tobacco (Nicotiana tabacum), and soybean (Glycine max). Transmission The transmission of EuMV has not been investigated. It is likely that, in common with other begomoviruses, EuMV is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. The virus was shown experimentally to infect E. heterophylla, Nicotiana benthamiana, Arabidopsis thaliana, Datura stramonium, bean (Phaseolus vulgaris) and tomato (Solanum lycopersicum) by biolistic inoculation with cloned virus genome components (Hernandez-Zepeda et al. 2007; Gregorio-Jorge et al. 2010). Infected E. heterophylla plants exhibit a golden mosaic and leaf malformation symptoms. Virion properties and genome The structure of EuMV particles has not been investigated. In common with all geminiviruses, the virions of EuMV are likely geminate (twinned icosahedra). EuMV is typical of the majority of begomoviruses native to the New World with a bipartite genome consisting of two circular, single-stranded DNA components. DNA-A contains 2613 nt (DQ318937 = NC_008304) and DNA-B of 2602 nt (DQ318938 = NC_008305) (Briddon 2001; Gregorio-Jorge et al. 2010; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the New World, the DNA-A component of EuMV encodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for EuMV.

Euphorbia heterophylla (Painted Euphorbia)

Euphorbia yellow mosaic virus Taxonomic position Genus: Begomovirus

977

(EuYMV)

Family: Geminiviridae

Geographical distribution EuYMV infection in plants of Euphorbia heterophylla was reported from Brazil (Fernandes et al. 2011). Some isolates of the virus were previously known as Euphorbia mosaic Peru virus. Symptoms and host(s) The virus-infected painted euphorbia plants exhibit bright yellow mosaic symptoms with some leaf crumpling. The virus has been isolated from Euphorbia heterophylla, Sida santaremensis, Macroptilium atropurpureum, tomato (Solanum lycopersicum), and Crotalaria juncea. Transmission The virus is transmitted by the vector of begomoviruses, the whitefly Bemisia tabaci (Mar et al. 2017). Although the mechanism of transmission has not been investigated, this will likely be in a circulative, non-propagative manner, in common with other begomoviruses which have been investigated. Experimentally the virus has been transmissible to Euphorbia heterophylla, pepper (Capsicum annuum), Datura stramonium, and Nicotiana benthamiana by biolistic inoculation of cloned virus components. Virion properties and genome The structure of EuYMV particles has not been investigated. In common with all geminiviruses, the virions of EuYMV are likely geminate (twinned icosahedra). EuYMV is typical of the majority of begomoviruses native to the New World with a bipartite genome consisting of two circular, single-stranded DNA components. DNA-A contains 2609 nt (FJ619507 = NC_012553; JF756676) and DNA-B of 2578 nt (FJ619508 = NC_012554) (Briddon 2001; Fernandes et al. 2011; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the New World, the DNA-A component of EuMV encodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for EuMV.

Merremia mosaic virus Taxonomic position Genus: Begomovirus

(MerMV)

Family: Geminiviridae

MerMV infection in plants of Euphorbia heterophylla was reported from Belize (McLaughlin et al. 2008). The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of MerMV, refer to Merremia spp.

Tomato severe rugose virus Taxonomic position Genus: Begomovirus

(ToSRV)

Family: Geminiviridae

E

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Euphorbia marginata (Snow-on-the-mountain)

ToSRV infection in plants of Euphorbia heterophylla was reported from Brazil (Barreto et al. 2013). The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of ToSRV, refer to Solanum lycopersicum.

References Barreto SS, Hallwass M, Aquino OM, Inoue-Nagata AK (2013) A study of weeds as potential inoculum sources for a tomato-infecting begomovirus in central Brazil. Phytopathology 103(5):436–444 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55. Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 CABI/EPPO (2009) Euphorbia mosaic virus. [Distribution map]. Distribution Maps of Plant Diseases, No.October, 1st edn. CABI, Wallingford. Map 1061 Collins AM, Brown JK, Rehman MM, Roye ME (2009) Complete nucleotide sequence of an isolate of Euphorbia mosaic virus that infects Euphorbia heterophylla and Wissadula amplissima in Jamaica. Arch Virol 154:1859–1860 Costa AS, Bennett CW (1950) Whitefly-transmitted mosaic of Euphorbia prunifolia. Phytopathology 40:266–283 Debrot E, Centeno F (1985) Occurrence of euphorbia mosaic virus infecting Euphorbia heterophylla L. in Venezuela. Agron Trop 35(4–6):5–12 Fernandes FR, Albuquerque LC, de Oliveira CL, Cruz ARR, da Rocha WB, Pereira TG, Naito FYB, Dias NM, Nagata T, Faria JC, Zerbini FM, Aragao FJL, Inoue-Nagata AK (2011) Molecular and biological characterization of a new Brazilian begomovirus, Euphorbia yellow mosaic virus (EuYMV), infecting Euphorbia heterophylla plants. Arch Virol 156:2063–2069 Fiallo-Olive E, Navas-Castillo J, Moriones E, Martínez-Zubiaur Y (2012) Begomoviruses infecting weeds in Cuba: increased host range and a novel virus infecting Sida rhombifolia. Arch. Virol. 157:141–146 Gregorio-Jorge J, Bernal-Alcocer A, Ban~uelos-Herna’ndez B, Alpuche-Solı’s AG, Herna’ndez-Zepeda C, MorenoValenzuela O, Frı’as-Trevin~o G, Argu¨ello-Astorga GR (2010) Analysis of a new strain of Euphorbia mosaic virus with distinct replication specificity unveils a lineage of begomoviruses with short Rep sequences in the DNA-B intergenic region. Virol J 7:275–290 Hernandez-Zepeda C, Idris AM, Carnevali G, Brown JK, Moreno-Valenzuela OA (2007) Molecular characterization and experimental host range of Euphorbia mosaic virus-Yucatan Peninsula, a begomovirus species in the Squash leaf curl virus clade. Plant Pathol 56:763–777 Mar TB, Mendes IR, Lau D, Fiallo-Olivé E, Navas-Castillo J, Alves MS, Murilo Zerbini F (2017) Interaction between the New World begomovirus Euphorbia yellow mosaic virus and its associated alphasatellite: effects on infection and transmission by the whitefly Bemisia tabaci. J Gen Virol 98(6):1552–1562 McLaughlin PD, McLaughlin WA, Maxwell DP, Roye ME (2008) Identification of begomoviruses infecting crops and weeds in Belize. Plant Viruses 2:58–63 Shepherd DN, Martin DP, Lefeuvre P, Monjane AL, Owor BE, Rybicki EP, Varsani A (2008) A protocol for the rapid isolation of full geminivirus genomes from dried plant tissue. J Virol Meth 149:97–102 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. Journal of General Virology 98:131–133

Euphorbia marginata (Snow-on-the-mountain) Family: Euphorbiaceae

Ornamental

Asclepias asymptomatic virus

(AsAV)

Taxonomic position AsAV is a tentative member of the genus Tymovirus and family Tymoviridae.

Euphorbia milii x lophogona

979

AsAV infection in plants of Euphorbia marginata was reported from Oklahoma (USA) (Hackett et al. 2009). The virus-infected Euphorbia marginata plants do not exhibit noticeable symptoms. The virus is transmitted by the beetle vectors, Tetraopes spp., and is also transmissible by mechanical sapinoculation. For more details of AsAV, refer to Asclepias spp.

References Hackett J, Muthukumar V, Wiley GB, Roe BA, Melcher U (2009) Viruses in Oklahoma Euphorbia marginata (Pursh). Proc Okla Acad Sci 89:57–62

Euphorbia milii x lophogona Family: Euphorbiaceae

Ornamental

Euphorbia ringspot virus Taxonomic position Genus: Potyvirus

(EuRSV)

Family: Potyviridae

Geographical distribution EuRSV infection in plants of Euphorbia milii x lophogona was reported from Germany, Denmark, Venezuela, India, and the USA (Bode and Lesemann 1976; Guaragna et al. 2004; Marys and Romano 2011; Jordan et al. 2011). Symptoms and host(s) The virus-infected Euphorbia milii x lophogona plants exhibit ring-shaped chlorotic spots on leaves, leaf and flower breaking, and growth reduction. Transmission The virus is transmitted by an aphid vector Myzus persicae in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to some members of the Euphorbiaceae and Chenopodiaceae. The virus is graft-transmissible and mechanically transmissible by cutting first diseased and then healthy stems with the same knife (Bode and Lesemann 1976). Virion properties and genome The virions are non-enveloped, filaments, and flexuous with a clear modal length of 750 nm and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA. A partial genome sequence of 2147 nt is available (AY697300) in addition to the full genome of 10,154 nt (KX355613 = NC_031339) (Bode and Lesemann 1976; Guaragna et al. 2004; Revers and Garcia 2015; Knierim et al. 2017; Wylie et al. 2017).

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Euphorbia pulcherrima (Poinsettia)

References Bode O, Lesemann DE (1976) Euphorbia ringspot virus, a new virus in Euphorbia milii X lophogona. Acta Hortic (ISHS) 59:161–166 Guaragna MA, Ambrose J, Jordan RL (2004) Detection and characterization of Euphorbia ringspot potyvirus. Phytopathology 94:S36 Jordan R, Guaragna MA, Putnam M (2011) Detection and molecular characterization of new and emerging potyviruses of ornamental plants. Acta Hortic (ISHS) 901:159–166 Knierim D, Menzel W, Winter S (2017) Analysis of the complete genome sequence of Euphorbia ringspot virus, an atypical member of the genus Potyvirus. Arch Virol 162:291–293 Marys E, Romano M (2011) Occurrence of Euphorbia ringspot virus in Euphorbia milii cv. splendens in Venezuela. J Phytopathol 159:66–68 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. Journal of General Virology 98:352–354

Euphorbia pulcherrima (Poinsettia) Family: Euphorbiaceae

Ornamental

Ageratum yellow vein virus Taxonomic position Genus: Begomovirus

(AYVV)

Family: Geminiviridae

AYVV infection in plants of Euphorbia pulcherrima was reported from China (Zhang et al. 2014). The virus-infected poinsettia plants exhibit leaf curl and vein thickening symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The virus is not transmissible by mechanical sap-inoculation nor by contact between plants. For more details of AYVV, refer to Ageratum spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV was reported infecting plants of Euphorbia pulcherrima in Iran (Ghotbi and Shahraeen 2012). The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Euphorbia leaf curl Guangxi virus Taxonomic position Genus: Begomovirus

(EuLCGxV)

Family: Geminiviridae

Euphorbia pulcherrima (Poinsettia)

981

Geographical distribution EuLCGxV infection in plants of Euphorbia pulcherrima was reported from China (Xu and Zhou 2007). Symptoms and host(s) The virus-infected poinsettia plants exhibit leaf curl and vein thickening symptoms. Transmission The virus has been shown to be transmitted by the whitefly Bemisia tabaci. Although the mechanism of transmission has not been investigated, this will likely be in a circulative, non-propagative manner, in common with other begomoviruses which have been investigated. Experimentally the virus has been transmissible to tobacco (Nicotiana tabacum), tomato (Solanum lycopersicum), Datura stramonium, and E. pulcherrima by insect transmission. Virion properties and genome The structure of the virions of EuLCGxV has not been investigated. In common with all geminiviruses, the virions of EuLCGxV are likely geminate (twinned quasi-icosahedra). EuLCGxV is a typical Old World monopartite begomovirus with a genome consisting of a single circular molecule of single-stranded DNA of 2747 nt (AM411424) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). The characterized genome of EuLCGxV encodes the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated. It is unclear whether the virus associates with a betasatellite.

Euphorbia leaf curl virus Taxonomic position Genus: Begomovirus

(EuLCuV)

Family: Geminiviridae

Geographical distribution EuLCuV infection in plants of Euphorbia pulcherrima was reported from China (Ma et al. 2004). Symptoms and host(s) The virus-infected poinsettia plants exhibit leaf curl and vein thickening symptoms. The virus was isolated from Euphorbia pulcherrima and passion fruit (Passiflora edulis  Passiflora edulis f. flavicarpa). Transmission The virus has been shown to be transmitted by the whitefly Bemisia tabaci. Although the mechanism of transmission has not been investigated, this will likely be in a circulative, non-propagative manner, in common with other begomoviruses which have been investigated. Experimentally, the virus has been transmissible to Nicotiana benthamiana, N. glutinosa, tomato (Solanum lycopersicum), Petunia hybrida, and tobacco (N. tabacum) by Agrobacterium-mediated inoculation of virus clones and whitefly transmission (Wu et al. 2011). Infected E. pulcherrima plants exhibit leaf curl and vein thickening symptoms. Virion properties and genome The structure of the virions of EuLCuV has not been investigated. In common with all geminiviruses, the virions of EuLCuV are likely geminate (twinned quasi-icosahedra).

E

982

Euphorbia pulcherrima (Poinsettia)

EuLCuV is a typical Old World monopartite begomovirus with a genome consisting of a single circular molecule of single-stranded DNA of 2746 nt (AJ558121 = NC_005319; FJ487911, KC161185, KC852148) (Briddon 2001; Ma et al. 2004; Brown et al. 2015; Zerbini et al. 2017). The genome of EuLCuV encodes the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated. Although the virus has been shown to be capable of interacting with a betasatellite (Wu et al. 2011), there are no reports of an association of the virus with a betasatellite in the field.

Euphorbia yellow leaf curl virus Taxonomic position Genus: Begomovirus

(EuYLCV)

Family: Geminiviridae

Geographical distribution EuYLCV infection in plants of Euphorbia pulcherrima was reported from Pakistan (Riaz and Ashfaq unpublished; KT159766). Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm, with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2731 nt (KT159766 = NC_038976) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Papaya leaf curl Guandong virus Taxonomic position Genus: Begomovirus

(PaLCuGdV)

Family: Geminiviridae

PaLCuGdV infection in plants of Euphorbia pulcherrima was reported from China (Zhang et al. 2014). The virus-infected poinsettia plants exhibit leaf curl and vein thickening symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The virus is not mechanically sap-transmissible. For more details of PaLCuGdV, refer to Carica papaya.

Poinsettia latent virus

(PnLV)

Synonyms Poinsettia cryptic virus (PnCV) Taxonomic position Genus: Polemovirus

Family: Solemoviridae

Euphorbia pulcherrima (Poinsettia)

983

Geographical distribution PnLV infection in plants of Euphorbia pulcherrima was first reported from Germany (Koenig and Lesemann 1980). The virus was also reported from Denmark (Paludan and Begtrup 1986). Symptoms and host(s) Virus-infected poinsettia plants do not exhibit any symptoms. Transmission Vegetative propagation and grafting are the main sources for virus dissemination. Virion properties and genome The virions are icosahedra of 34 nm in diameter and have a T = 3 icosahedral symmetry. The genome of PnLV is a single molecule of linear positive-sense ssRNA of 4652 nt (AJ867490 = NC_011543); it has a 50 VPg. Particles may also contain sgRNAs of 1.1 and 2.6 kb. PnLV genome consists of four ORFs (Aus dem Siepen et al. 2005; Somera et al. 2015).

Poinsettia mosaic virus Taxonomic position Genus: Unassigned

(PnMV)

Family: Tymoviridae

Geographical distribution PnMV was first reported in plants of Euphorbia pulcherrima by Fulton and Fulton (1980) from Wisconsin, USA. The virus is worldwide in distribution, including Australia, Canada, the USA, Germany, Finland, Denmark, New Zealand, and the UK (Koenig and Lesemann 1980; Chiko 1983; Guy 1985; Paludan and Begtrup 1986; Bertaccini et al. 1996; Carballo et al. 2001; Lebas et al. 2007; Lemmety et al. 2008; Fisher and Bell 2013; Ocampo et al. 2013). Symptoms and host(s) The virus-infected poinsettia plants exhibit systemic mosaic and malformation symptoms; bracteoles are malformed and may not develop their full cream or red coloration (Pfannenstiel et al. 1982). The natural host range of this virus is restricted to poinsettia (Euphorbia pulcherrima). Transmission No vector is reported for transmission of this virus. The virus is mechanically sap-transmissible and not transmissible by contact between plants. Graft-transmission is recorded, but no seed and pollen transmission (Floeistad and Blystad 1999; Blystad and Floeistad 2002). The virus is transmitted by the use of virus-infected propagative material. Virion properties and genome The virions are isometric, approximately 35 nm in diameter (Gordon et al. 1996). The genome is a single molecule of positive-sense single-stranded RNA of 6099 nt (AJ271595 = NC_002164), and the capsid is composed of 21.7-kDa protein subunits (Pfannensteil et al. 1982; Lesemann et al. 1983; Bradel et al. 2000; Martelli et al. 2002).

E

984

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

Euphorbia pulcherrima (Poinsettia)

(TMV)

Family: Virgaviridae

TMV infection in plants of Euphorbia pulcherrima was reported from Mexico (Ocampo et al. 2013). The virus-infected poinsettia plants exhibit symptoms of chlorosis, mosaic, leaf distortion, and white spots. There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

References Aus dem Siepen MS, Pohl JO, Koo BJ, Wege C, Jeske H (2005) Poinsettia latent virus is not a cryptic virus, but a natural polerovirus-sobemovirus hybrid. Virology 336:240–250 Bertaccini A, Vibio M, Bellardi MG (1996) Virus diseases of ornamental shrubs. X. Euphorbia pulcherrima Willd. infected by viruses and phytoplasma. Phytopath Medit 35:129–132 Blystad DR, Floeistad E (2002) Transmission of Poinsettia mosaic virus. Acta Hortic 568:225–226 Bradel BG, Preil W, Jeske H (2000) Sequence analysis and genome organization of Poinsettia mosaic virus (PnMV) reveal closer relationship to marafiviruses than to tymoviruses. Virology 271:289–297 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Carballo O, Izaguirre ML, Marys E (2001) Detection of Poinsettia mosaic virus infecting Poinsettias (Euphorbia pulcherrima) in Venezuela. Plant Dis 85:1208 Chiko AW (1983) Poinsettia mosaic virus in British Columbia. Plant Dis 67:427–428 Fisher JR, Bell J (2013) First report of Poinsettia mosaic virus associated with a virus-like mottle symptom on poinsettia in Ohio. Plant Health Progress. https://doi.org/10.1094/PHP-2013-0813-01-BR. Online Floeistad E, Blystad DR (1999) Two new hosts for Poinsettia mosaic virus. Plant Dis 83:399 Fulton RW, Fulton JL (1980) Characterization of a tymo-like virus common in poinsettia. Phytopathology 70:321–324 Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses affecting propagated ornamentals in Northern Iran. Int Res J Microbiol 3:373–381 Gordon D, Morrison BC, Teakle DS (1996) Poinsettia mosaic virus – first recorded in Queensland. Aust Plant Pathol 25:145 Guy P (1985) New plant disease record in Tasmania: Poinsettia mosaic virus. Aust Plant Pathol 14:12–13 Koenig R, Lesemann D-E (1980) Two isometric viruses in poinsettias. Plant Dis 64:782–784 Lebas BSM, Ochoa-Corona FM, Elliott DR, Tang JZ, Alexander BJR (2007) Detection of Poinsettia mosaic virus by RT-PCR in Euphorbia spp. in New Zealand. Plant Dis 91:110 Lemmety A, Laamanen J, Soukainen M, Tegel J (2008) Emerging virus and viroid pathogen species identified for the first time in horticultural plants in Finland in 1997–2010. Agric Food Sci 20:29–41 Lesemann D-E, Koenig R, Huth W, Brunt AA, Phillips S, Barton RJ (1983) Poinsettia mosaic virus: a tymovirus? Phytopath Z 107:250–262 Ma X, Cai J, Li G, Qin B, Zhou X (2004) Molecular characterization of a distinct begomovirus infecting Euphorbia pulcherrima in China. J Phytopathol 152:215–218 Martelli GP, Sabanadzovic S, Abou-Ghanem Sabanadzovic N, Edwards MC, Dreher T (2002) The family Tymoviridae. Arch Virol 147(9):1837–1846 Ocampo T, Ochoa Martinez DL, Ramirez Rojas S, Valdovinos Ponce G, Nava DC (2013) First report of Tobacco mosaic virus (TMV) and Poinsettia mosaic virus (PnMV) in poinsettia (Euphorbia pulcherrima Willd. ex Klotzsch) in Mexico. Phyton (Buenos Aires) 82:235–241 Paludan N, Begtrup J (1986) Inactivation of Poinsettia mosaic virus and Poinsettia cryptic virus in Euphorbia pulcherrima using heat treated mini-cuttings and meristem-tip culture. Tiddsskr Planteavl 90:283–290 Pfannenstiel MA, Mintz KP, Fulton RW (1982) Evaluation of heat therapy of poinsettia mosaic and characterization of the viral components. Phytopathology 72:252–254 Somera M, Sarmiento C, Truve E (2015) Overview on Sobemoviruses and a Proposal for the Creation of the Family Sobemoviridae. Viruses 7:3076–3115

Eustoma russellianum (Lisianthus; Russell prairie gentian)

985

Wu J, Zulfiqar A, Huang C (2011) Infectivity of Euphorbia leaf curl virus and interaction with Tomato yellow leaf curl China betasatellite. Arch Virol 156(3):517–521 Xu YP, Zhou XP (2007) A new begomovirus associated with leaf curl disease of Euphorbia pulcherrima. J Plant Pathol 89:S69 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. Journal of General Virology 98:131–133 Zhang J, Cui GJ, Yang CX, Wu ZJ (2014) First report of Ageratum yellow vein virus and Papaya leaf curl Guangdong virus on Euphorbia pulcherrima in China. J Plant Pathol 96:S4.128

Eustoma russellianum (Lisianthus; Russell prairie gentian) Synonyms Eustoma grandiflorum Family: Gentianaceae

Ornamental

Ageratum yellow vein virus Taxonomic position Genus: Begomovirus

(AYVV)

Family: Geminiviridae

AYVV infection in plants of Eustoma russellianum was reported from Taiwan (Cheng et al. 2005). The virus-infected lisianthus plants exhibit foliar vein-clearing symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The virus is not transmissible by either mechanical inoculation or contact between plants. For more details of AYVV, refer to Ageratum spp.

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

ArMV infection in plants of Eustoma spp. was reported from Iran (Ghotbi and Shahraeen 2005). The virus-infected lisianthus plants exhibit symptoms of mosaic, leaf chlorosis, small necrotic lesions, and leaf malformation and deformation. This virus is transmitted by the nematode vectors (Xiphinema spp.) in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

E

986

Eustoma russellianum (Lisianthus; Russell prairie gentian)

BYMV infection in plants of Eustoma grandiflorum was first reported from Northern Italy in 1984 (Lisa and Dellavalle 1987). The virus spreads in Israel and Taiwan (Chang and Lin 1990; Gera and Cohen 1990; Chang and Tsai 1993). The virus-infected lisianthus plants exhibit symptoms of pronounced stunting, leaf curling, mosaic, chlorotic spotting, and color break in flowers. The virus is transmitted by a number of aphids in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Bidens mottle virus Taxonomic position Genus: Potyvirus

(BiMoV)

Family: Potyviridae

BiMoV infection in plants of Eustoma russellianum was reported from Taiwan (Chen et al. 2016a). The virus-infected lisianthus plants exhibit conspicuous systemic foliar chlorosis with solid if not hollow (sunken) spot symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BiMoV, refer to Bidens spp.

Broad bean wilt virus Taxonomic position Genus: Fabavirus

(BBWV)

Family: Secoviridae

BBWV (but without specifying either of the currently recognized species BBWV-1 or BBWV-2) infection in plants of Eustoma russellianum was reported from Japan and Taiwan (Iwaki et al. 1985; Yamashita 1990; Chen et al. 2002). The virus-infected lisianthus plants showed yellow spots, with irregular concentric rings of pale and dark green tissue, becoming necrotic as leaves aged; oak leaf line patterns and terminal necrosis have also been reported. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BBWV, refer to Vicia faba.

Carnation mottle virus Taxonomic position Genus: Alphacarmovirus

(CarMV)

Family: Tombusviridae

CarMV infection in plants of Eustoma grandiflorum was reported from Taiwan (Chen et al. 2011). The virus-infected lisianthus plants exhibit symptoms of necrotic spots on leaves. The virus is transmitted by means not involving a vector. The virus is transmissible by mechanical sap-inoculation, by grafting, and also by contact between plants. For more details of CarMV, refer to Dianthus caryophyllus.

Chrysanthemum stem necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(CSNV)

Eustoma russellianum (Lisianthus; Russell prairie gentian)

987

CSNV infection in plants of Eustoma grandiflorum was reported from Japan and Brazil (Momonoi et al. 2011; Duarte et al. 2014). The virus-infected lisianthus plants exhibit symptoms of necrotic streaks on stems and necrosis on leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of CSNV, refer to Chrysanthemum spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

E Family: Bromoviridae

CMV is reported in plants of Eustoma grandiflorum from the USA, Taiwan, Italy, Iran, and Israel (Provvidenti 1985; Gera and Cohen 1990; Gera 1994; Chen and Hu 1999; Ghotbi and Shahraeen 2004). The virus-infected lisianthus plants exhibit mosaic and distortion of leaves and stunting; flowers showed color break and were malformed. A CMV isolate from Italy has exhibited milder symptoms like chlorotic mottle on leaves and slight reduction in plant growth. The virus is transmitted by an aphid vector Aphis gossypii in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Eggplant mottled crinkle virus

(EMCV)

Synonyms Lisianthus necrosis virus (LNV)

Taxonomic position Genus: Tombusvirus

Family: Tombusviridae

LNV was first identified in plants of Eustoma grandiflorum in Japan (Iwaki et al. 1987) and later detected in Taiwan (Chen et al. 2000, 2001, 2006). The virus-infected lisianthus plants show systemic bright yellow chlorosis and ringspots followed by necrosis. Flowers exhibit white stripes. The virus is mechanically sap-transmissible. No vector is reported for this virus. For more details of EMCV, refer to Solanum melongena.

Groundnut ringspot orthotospovirus Taxonomic position Genus: Orthotospovirus

(GRSV)

Family: Tospoviridae

GRSV infection in plants of Eustoma grandiflorum was reported from Brazil (Alexandre et al. 1999, 2004). The virus-infected lisianthus plants show necrotic spots and line pattern symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of GRSV, refer to Arachis hypogaea.

988

Eustoma russellianum (Lisianthus; Russell prairie gentian)

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Eustoma grandiflorum was reported from the USA, Japan, and Hungary (Daughtrey 1996; McGovern et al. 1997; Doi and Kato 2003; Uga and Tsuda 2005; Toth et al. 2007). The virus-infected lisianthus plants exhibit necrotic ringspots, leaf distortion, systemic necrosis, wilting, stunting, and death of the plants. The virus is transmitted by a thrips vector Frankliniella occidentalis in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

IYSV was reported in plants of Eustoma grandiflorum in Israel, Japan, the Netherlands, and the UK (Kritzman et al. 2000, 2001; Gera et al. 2002; Doi et al. 2003; Anon 2007; Mumford et al. 2008; Fuji et al. 2015). The virus-infected lisianthus plants were stunted, had necrotic spots and rings on leaves and stems, and developed tip necrosis and flower distortion (Kritzman et al. 2000). The virus is transmitted by the onion thrips (Thrips tabaci) in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of IYSV, refer to Iris spp. or Allium cepa.

Lettuce mosaic virus Taxonomic position Genus: Potyvirus

(LMV)

Family: Potyviridae

LMV was reported in plants of Eustoma grandiflorum in Italy (Lisa and Gera 1995; Lisa et al. 1995). The virus-infected lisianthus plants show yellow mosaic on leaves, color breaking, and malformation of flowers along with reduced growth of the plants. The virus is transmitted by aphid vectors in a nonpersistent manner, and is also transmissible by mechanical sap-inoculation. For more details of LMV, refer to Lactuca sativa.

Lisianthus enation leaf curl virus Taxonomic position Genus: Begomovirus

(LELCV)

Family: Geminiviridae

Geographical distribution LELCV infection in plants of Eustoma grandiflorum was reported from Taiwan (Chen and Chao, unpublished – LC091538)

Eustoma russellianum (Lisianthus; Russell prairie gentian)

989

Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm, with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2759 nt (LC091538 = NC_031466) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Lisianthus necrotic ringspot virus

(LNRV)

Taxonomic position LNRV is a tentative member of the genus Orthotospovirus and family Tospoviridae Geographic distribution LNRV infection in plants of Eustoma grandiflorum was reported from Japan (Shimomoto et al. 2014). Symptoms and host(s) The virus-infected plants exhibit necrotic ringspot symptoms. LNRV is only known to naturally infect lisianthus (Eustoma grandiflorum), experimental inoculations suggest a very restricted host range. Transmission The natural vector of this virus is unknown, but is presumed to be a species of thrips. The virus is not transmitted by either Frankliniella occidentalis or Thrips tabaci, the most prevalent thrips in Japanese greenhouses (Shimomoto et al. 2014). The virus is mechanically sap-transmissible. Virion properties and genome Quasi-isometric particles of c.160 nm diameter were observed. The full genome includes three negative or ambisense single-stranded RNAs (the L, M, and S RNAs, in order of decreasing size); the full sequence of only the S RNA segment of 2768 nt has been determined (AB852525) (Shimomoto et al. 2014).

Moroccan pepper virus Taxonomic position Genus: Tombusvirus

(MPV)

Family: Tombusviridae

MPV infection in plants of Eustoma grandiflorum was reported from Iran and Japan (Beikzadeh et al. 2011; Ohki et al. 2014). The virus-infected lisianthus plants produce yellow and necrotic spots on the leaves and became severely deformed because of a strong leaf curling and the production of shorter internodes. The virus is transmissible by mechanical inoculation to 3–9 families. The virus is transmissible by grafting. No vector is known for this virus. For more details of MPV, refer to Capsicum annuum.

E

990

Eustoma russellianum (Lisianthus; Russell prairie gentian)

Papaya leaf curl Guandong virus Taxonomic position Genus: Begomovirus

(PaLCuGdV)

Family: Geminiviridae

PaLCuGdV infection in plants of Eustoma grandiflorum was reported from Taiwan (Chen et al. 2016b). The virus-infected lisianthus plants exhibit leaf curl, cup-shaped upper leaves, and enations both on the cup-shaped upper leaves and on flower petals. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The virus is not mechanically sap-transmissible. For more details of PaLCuGdV, refer to Carica papaya.

Pepper ringspot virus Taxonomic position Genus: Tobravirus

(PepRSV)

Family: Virgaviridae

PepRSV infection in plants of Eustoma grandiflorum was reported from Brazil (Rivas et al. 2000). The virus-infected lisianthus plants exhibit ringspot symptoms. The virus is transmitted by nematode vectors and is also transmissible by mechanical sap-inoculation. For more details of PepRSV, refer to Capsicum annuum.

Pepper veinal mottle virus Taxonomic position Genus: Potyvirus

(PVMV)

Family: Potyviridae

PVMV infection in plants of Eustoma grandiflorum was reported from Taiwan (Cheng et al. 2009). The virus-infected lisianthus plants exhibit stunting and yellow spotting symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants. For more details of PVMV, refer to Capsicum annuum.

Pothos latent virus Taxonomic position Genus: Aureusvirus

(PoLV)

Family: Tombusviridae

PoLV infection in plants of Eustoma grandiflorum was reported from Taiwan (Chen et al. 2016c). The virus-infected lisianthus plants exhibit necrotic ringspots and concentric line pattern symptoms. The virus is transmitted through soil and water without a vector, and is also transmissible by mechanical sapinoculation to herbaceous plants. For more details of PoLV, refer to Scindapsus spp.

Eustoma russellianum (Lisianthus; Russell prairie gentian)

Sweet potato chlorotic stunt virus

991

(SPCSV)

Synonyms Sweet potato sunken vein virus (SPSVV) Taxonomic position Genus: Crinivirus

Family: Closteroviridae

SPSVV infection in plants of Eustoma grandiflorum was reported from Israel (Cohen et al. 2001). The virus-infected lisianthus plants show no obvious symptoms except stunting of flower stems. The virus is transmitted by the whitefly vector Bemisia argentifolii in a semi-persistent manner. The virus is not mechanically sap-transmissible. For more details of SPCSV, refer to Ipomoea batatas.

Tobacco mosaic virus

(TMV)

Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

TMV was reported in plants of Eustoma grandiflorum from Israel (Gera and Cohen 1990). The virusinfected lisianthus plants show typical mosaic symptoms. The virus is highly infectious and mechanically sap-transmissible to a large number of hosts and also by contact between plants. There is no known vector is reported for this virus. For more details of TMV, refer to Nicotiana tabacum.

Tobacco streak virus

(TSV)

Synonyms Lisianthus line pattern virus Taxonomic position Genus: Ilarvirus

Family: Bromoviridae

TSV infection in plants of Eustoma grandiflorum has been reported from Brazil (earlier Lisianthus line pattern virus was identified in Italy as a possible strain of TSV) (Lisa et al. 1994). The virus spreads in Italy, Israel, and Brazil (Lisa and Gera 1995; de Freitas et al. 1996). The virus is specific to lisianthus on which it causes necrotic spots and line patterns on leaves and without symptoms on flowers (Lisa and Gera 1995). The virus is transmitted by the thrips vectors, the virus present in/on pollen, enter the host through injuries caused by thrips while feeding. The virus is also transmissible by mechanical sapinoculation. The virus is not transmissible by contact between plants (Lisa and Gera 1995). For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato bushy stunt virus Taxonomic position Genus: Tombusvirus

(TBSV)

Family: Tombusviridae

E

992

Eustoma russellianum (Lisianthus; Russell prairie gentian)

TBSV infection in plants of Eustoma grandiflorum was reported from Japan (Fujinaga et al. 2006). The virus-infected lisianthus plants exhibit necrotic spotting and stunting symptoms. The virus is transmissible by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants. No vector is known for this virus. For more details of TBSV, refer to Solanum lycopersicum.

Tomato chlorotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(TCSV)

Family: Tospoviridae

TCSV infection in plants of Eustoma grandiflorum was reported from Argentina and Brazil (Alexandre et al. 1999; Dal Bo et al. 1999, 2001; Gracia et al. 1999). The virus-infected lisianthus plants exhibit necrotic spots, rings, and line pattern symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of TCSV, refer to Solanum lycopersicum.

Tomato mosaic virus Taxonomic position Genus: Tobamovirus

(ToMV)

Family: Virgaviridae

ToMV infection in plants of Eustoma grandiflorum was reported from Italy, Argentina, and Taiwan (Wolcan et al. 1996; Jan et al. 2003; Zheng et al. 2003). The virus-infected lisianthus plants exhibit mosaic and necrotic/chlorotic ring patterns on leaves and whitish stripes on flowers, with stunting and light brown necrotic rings and spots on the lower leaves (Lisa and Dellavalle 1989). No vector is involved in the spread of this virus. This virus is transmissible by mechanical sap-inoculation by grafting, and by contact between plants. For more details of ToMV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Eustoma grandiflorum was reported from the USA, Israel, Italy, Argentina, Korea, and Japan (Lisa et al. 1990; Hausbeck et al. 1992; Wolcan et al. 1996; Kuroda et al. 1999; Yoon et al. 2017). The virus-infected lisianthus plants show symptoms of yellowing, mosaic, and necrosis on leaves, and color breaking was noticed on flowers (Kuroda et al. 1999). The virus is transmitted by thrips vectors in a persistent-propagative manner. The virus is transmissible by mechanical sapinoculation to a large number of herbaceous hosts and by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Eustoma russellianum (Lisianthus; Russell prairie gentian)

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

993

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Eustoma grandiflorum was reported from Israel and Korea (Cohen et al. 1993, 1995; Kil et al. 2014). The virus-infected lisianthus plants exhibit leaf curling and stunting symptoms. The growing apex of the plant is distorted, and the leaves were cup shaped; typical enations are noticed on the lower surface of the leaves, and the plants were stunted (Kil et al. 2014). The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The virus is transmissible by grafting but not transmissible by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Eustoma grandiflorum was reported from Taiwan (Chao et al. 1998, 2000). The virus-infected lisianthus plants exhibit systemic yellow spotting and stunting symptoms. The virus is transmitted by the aphid vectors Myzus persicae and Brevicoryne brassicae in a non-persistent manner. The virus is also mechanically sap-transmissible and causes systemic mosaic symptoms on many cruciferous crops. For more details of TuMV, refer to Brassica rapa.

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV infection in plants of Eustoma grandiflorum was reported from Japan (Inoue and Kasuyama 2001). The virus-infected lisianthus plants exhibit chlorotic or whitish ringspot symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of WMV, refer to Citrullus lanatus.

Youcai mosaic virus Taxonomic position Genus: Tobamovirus

(YoMV)

Family: Virgaviridae

YoMV infection in plants of Eustoma grandiflorum was reported from Japan (Yamashita and Fukui 2002). There is no known vector is reported for this virus. The virus is mechanically sap-transmissible and is also transmissible through contact between plants. For more details of YoMV, refer to Brassica napus.

E

994

Eustoma russellianum (Lisianthus; Russell prairie gentian)

References Alexandre MAV, Duarte LML, Rivas EB, Chagas CM (1999) Mixed infection by Tospovirus in ornamental crops in São Paolo State, Brazil. Summa Phytopathol 25:353–356 Alexandre MAV, Seabra PV, Rivas EB, Duarte LML, Galleti SR (2004) Viruses, viroids, phytoplasmas and spiroplasmas detected in ornamental plants from 1992 to 2003. Rev Bras Hortic Ornamental 11:49–57 Anon (2007) Iris yellow spot virus detected in Eustoma in the Netherlands. EPPO reporting service 2007/008 Beikzadeh N, Peters D, Hassani-Mehraban A (2011) First report of Moroccan pepper virus on Lisianthus in Iran and worldwide. Plant Dis 95:1485 Briddon RW (2001) Begomovirus. Geminiviridae. The Springer index of plant viruses. Springer, New York, pp 340–348, https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A. (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Chang CA, Lin HH (1990) Isolation of Bean yellow mosaic virus from lisianthus developing foliar mosaic and flower breaking symptoms. Plant Pathol Bull 2:250–251 Chang CA, Tsai HT (1993) Isolation of Bean yellow mosaic virus from lisianthus developing foliar mosaic and flower breaking symptoms. Plant Pathol Bull 2:250–252 Chao CH, Chen CC, Chang CA (1998) Identification of turnip mosaic potyvirus isolated from Lisianthus. Plant Pathol Bull 7:223 Chao CH, Chen CC, Chang CA, Chen CC (2000) Identification of a Turnip mosaic virus isolate causing systemic yellow spotting on lisianthus. Plant Pathol Bull 9:115–122 Chen CC, Hu CC (1999) Purification and characterization of a cucumovirus from Lisianthus russellianum. Plant Prot Bull 41:179–198 Chen CC, Chen YK, Hsu HT (2000) Characterization of a virus infecting lisianthus. Plant Dis 84:506–509 Chen CC, Tsao SL, Hsu HT (2001) Diagnosis of Lisianthus necrosis virus infection by light and electron microscopy and serological assay. Plant Pathol Bull 10:105–114 Chen CC, Hu CC, Chen YK, Hsu HT (2002) A fabavirus inducing ringspot disease in lisianthus. Acta Hortic 568:51–57 Chen YK, Jan FJ, Chen CC, Hsu HT (2006) A new natural host of Lisianthus necrosis virus in Taiwan. Plant Dis 90:1112 Chen Y-K, Chang Y-S, Chen C-C (2011) Identification of carnation mottle virus from Lisianthus plants in Taiwan. Plant Dis 95:1036 Chen YK, Lee JY, Chang YS, Wu MY (2016a) First report of Bidens mottle virus causing chlorotic hollow spots on lisianthus in Taiwan. Plant Dis 100:1250–1251 Chen YK, Chao HY, Shi PJ, Tsai WY, Chao CH (2016b) First report of Papaya leaf curl Guangdong virus infecting lisianthus in Taiwan. Plant Dis 100:2342 Chen YK, Chang YS, Chao HY (2016c) Identification and characterization of Pothos latent virus causing necrotic ringspots and line patterns on lisianthus (Eustoma grandiflorum) in Taiwan. J Phytopathol 164(9):650–658 Cheng YH, Chen CC, Chang CA (2005) Whitefly-transmitted geminiviruses in ornamental plants and their control strategies in Taiwan. In: Proceedings of the international seminar on whitefly management and their control strategy. Food Fertil Technol Cent Agric Res Inst Pub Taichung, Taiwan ROC, p.95–105 Cheng YH, Chen CC, Liao JY, Deng TC, Chang CA (2009) Isolation and identification of pepper veinal mottle virus on lisianthus. J Taiwan Agric Res 58:196–207 Cohen J, Gera A, Gokkes M, Aker R, Antignus Y (1993) Tomato yellow leaf curl virus (TYLCV) is the causal agent of a new disease in lisianthus (Eustoma russellianum). Hassadeh 74:169–171 Cohen J, Gera A, Ecker R, Ben Joseph R, Perlsman M (1995) Lisianthus leaf curl a new disease of lisianthus caused by Tomato yellow leaf curl virus. Plant Dis 79:416–420 Cohen J, Lapidot M, Loebenstein G, Gera A (2001) First report of sweet potato sunken vein virus occurring in Lisianthus. Plant Dis 85:679 Dal Bo E, Chiarrone G, Rolleri J, Ronco L (1999) Tospovirus on ornamental crops in La Plata. Rev Fac Agron La Plata 104:35–40 Dal Bo E, Chiarrone G, Rolleri J, Ronc L (2001) New tospoviruses found in La Plata. Rev Fac Agron (La Plata) 104:35–40 Daughtrey ML (1996) Detection and identification of tospoviruses in greenhouses. Acta Hortic 431:90–98 de Freitas JC, Kitajima EW, Rezende JAM (1996) First report of tobacco streak virus on lisianthus in Brazil. Plant Dis 80:1080 Doi M, Kato K (2003) Necrotic spot disease of lisianthus caused by Impatiens necrotic spot virus (INSV). Annu Rep Kanto-Tosan Plant Prot Soc 50:83–87 Doi M, Zen S, Okuda M, Nakamura H, Kato K, Hanada K (2003) Leaf necrosis disease of lisianthus (Eustoma grandiflorum) caused by Iris yellow spot virus. Jpn J Phytopathol 69:181–188

Eustoma russellianum (Lisianthus; Russell prairie gentian)

995

Duarte LML, Vaz Alexandre MA, Gobatto D, Kitajima EW, Harakava R (2014) First report of Chrysanthemum stem necrosis virus on Russell prairie gentian in Brazil. Plant Dis 98:285 Fuji S, Zen S, Sato I, Kishi H, Furuya H, Okuda M (2015) Population dynamics of Iris yellow spot virus in onion fields and lisianthus greenhouses in Japan. Plant Pathol 64:808–816 Fujinaga M, Morikawa T, Doi M, Yoneyama C, Ibrahim M, Ogiso H, Miyamoto K, Miyasaka M, Ohki T, KameyaIwaki M, Natsuaki T (2006) Two tombusviruses isolated from lisianthus [Eustoma grandiflorum (Raf.) Shinn.] with necrotic stunt. Jpn J Phytopathol 72:109–115 Gera A (1994) The natural occurrence of Cucumber mosaic virus in ornamentals in Israel. Acta Hortic 377:99–106 Gera A, Cohen J (1990) The natural occurrence of bean yellow mosaic, cucumber mosaic, and tobacco mosaic viruses in lisianthus in Israel. Plant Pathol 39:561–564 Gera A, Kritzman A, Beckelman H, Cohen J, Raccah B (2002) Detection of Iris yellow spot virus in lisianthus. Acta Hortic (ISHS) 568:43–49 Ghotbi T, Shahraeen N (2004) Occurrence of cucumber mosaic virus (CMV) on Lisianthus from Varamin area. The First national festival and seminar on cut flowers. Tehran- Pakdasht 6–7 Oct. 62 Ghotbi T, Shahraeen N (2005) First report on incidence of Arabis mosaic virus (ArMV, Nepovirus) on ornamental plants in Iran. Iran J Plant Pathol 41(2):305–306 Gracia O, De Borbon CM, Millan D, Granval N, Cuesta GV (1999) Occurrence of different tospoviruses in vegetable crops in Argentina. J Phytopathol 147:223–227 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among green house ornamentals in Pennsylvania. Plant Dis 76:795–800 Inoue K, Kasuyama S (2001) Occurrence of ring spot of Russell prairie gentian [Eustoma grandiflorum] caused by watermelon mosaic virus, and detection of viruses of Russell prairie gentian in Okayama prefecture [Japan]. Bulletin of the Agricultural Experiment Station. Okayama Prefectural Gen Agric Cent (Jpn) 19:45–51 Iwaki M, Maria ERA, Hanada K, Onogi S, Zenbayashi R (1985) Three viruses occurred in lisianthus plants. Ann Phytopathol Soc Jpn 52:355 Iwaki M, Handa K, Maria ERA, Onogi S (1987) Lisianthus necrosis virus, a new necrovirus from Eustoma russellianumm. Phytopathology 77:867–870 Jan JF, Chen CC, Hsu HT (2003) Identification of Tomato mosaic virus infecting lisianthus in Taiwan. Plant Dis 87:1537 Kil EJ, Byun HS, Kim S, Hwang H, Kim MK, Kim CS, Choi HS, Lee KY, Lee S (2014) First report of Tomato yellow leaf curl virus infecting Eustoma (Eustoma grandiflorum) in Korea. Plant Dis 98:1163 Kritzman A, Beckelman H, Alexandrov S, Cohen J, Lampel M, Zaidan M, Raccah B, Gera A (2000) Lisianthus leaf necrosis: a new disease of lisianthus caused by Iris yellow spot virus. Plant Dis 84:1185–1189 Kritzman A, Beckelman H, Svetlana A, Cohen J, Miriam L, Raccah B, Gera A (2001) Detection of iris yellow spot virus in lisianthus. Phytoparasitica 29(3):256–257 Kuroda T, Katsube K, Suzuki K (1999) Spotted wilt disease of Prairie gentian (Eustoma grandiflorum (Raf.) Shinn.) caused by Tomato spotted wilt virus (TSWV). Jpn J Phytopathol 65:490–499 Lisa V, Dellavalle G (1987) Bean yellow mosaic virus in Lisianthus russelianum. Plant Pathol 36:214–215 Lisa V, Dellavalle G (1989) Malattie da virus del lisianthus. Giorn Studio Sul Lisianthus, Sanremo Lisa V, Gera A (1995) Lisianthus. In: Loebenstein G, Lawson RH, Brunt AA (eds) Virus and virus-like diseases of bulb and flower crops. Wiley, Chichester, pp 443–448 Lisa V, Vaira AM, Milne RG, Luisoni E, Rapetti S (1990) Tomato spotted wilt virus in five crops in Liguria. Inf Fitopatol 40:34–41 Lisa V, d’Aquilio M, Dellavalle G, Masenga V, Milne RG, Boccardo G (1994) Characterization of an ilarvirus from lisianthus (Eustoma grandiflorum). Acta Hortic 377:81–89 Lisa V, Dellavalle G, Ramasso E (1995) Lettuce mosaic potyvirus in lisianthus. Informatore Fitopatologico 45(3):58–59 McGovern RJ, Polston JE, Harbaugh BK (1997) Detection of a severe isolate of Impatiens necrotic spot virus infecting lisianthus in Florida. Plant Dis 81(11):1334 Momonoi K, Moriwaki J, Morikawa T (2011) Stem necrosis of aster and Russell prairie gentian caused by Chrysanthemum stem necrosis virus. J Gen Plant Pathol 77:142–146 Mumford RA, Glover R, Daly M, Nixon T, Harju V, Skelton A (2008) Iris yellow spot virus (IYSV) infecting lisianthus (Eustoma grandiflorum) in the UK: first finding and detection by real-time PCR. Plant Pathol 57:768 Ohki T, Kajihara H, Maoka T (2014) First report of Moroccan pepper virus on lisianthus (Eustoma grandiflorum) in Japan. J Gen Plant Pathol 80:90–93 Provvidenti R (1985) Two newly recognized hosts of cucumber mosaic virus: Eustoma grandiflorum and Peristrophe angustifolia. Plant Dis 69:542 Rivas EB, Galleti SR, Duarte LML, Seabra PV, Alexandre MAV (2000) Virus and phytoplasma diseases of lisianthus. Summa Phytopathol 26:257–262 Shimomoto Y, Kobayashi K, Okuda M (2014) Identification and characterization of Lisianthus necrotic ringspot virus, a novel distinct tospovirus species causing necrotic disease of lisianthus (Eustoma grandiflorum). J Gen Plant Pathol 80:169–175

E

996

Evolvulus spp.

Toth EK, Kriston E, Takacs A, Bajtek M, Kazinczi G, Horvath J (2007) First report of impatiens necrotic spot virus in ornamental plants in Hungary. Plant Dis 91:331 Uga H, Tsuda S (2005) A one-step reverse transcription-polymerase chain reaction system for the simultaneous detection and identification of multiple tospovirus infections. Phytopathology 95:166–171 Wolcan S, Ronco L, Bo ED, Lori G, Alippi H (1996) First report of diseases on lisianthus in Argentina. Plant Dis 80:223 Yamashita K (1990) Mosaic diseases occurred on gentian and prairie gentian in Aomori prefecture. Ann Rep Soc Plant Prot N Jpn 41:80–82 Yamashita K, Fukui Y (2002) A tobamovirus isolated from lisianthus is similar to Youcai mosaic virus (YMoV). Jpn J Phytopathol 68:235 Yoon JY, Choi GS, Choi SK (2017) First report of tomato spotted wilt virus in Eustoma grandiflorum in Korea. Plant Dis 101:515 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. Journal of General Virology 98:131–133 Zheng YX, Chen CC, Chao CH, Chang CA, Jan FJ (2003) Identification of Tomato mosaic virus infection in lisianthus in Taiwan. Plant Path Bul 12:290

Evolvulus spp. Family: Convolvulaceae

Tobacco streak virus Taxonomic position Genus: Ilarvirus

Ornamental

(TSV)

Family: Bromoviridae

According to Horst (2012), TSV has been reported from plants of Evolvulus spp. in the USA. The virus is transmitted by the thrips vectors, the virus present in/on pollen, enters the host through injuries by thrips while feeding. The virus is also mechanically sap-transmissible but is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

References Horst RK (2012) Westcott’s plant disease handbook, 5th edn. Springer, New York

Exacum spp. Family: Gentianaceae

Broad bean wilt virus Taxonomic position Genus: Fabavirus

Ornamental

(BBWV)

Family: Secoviridae

Exacum spp.

997

BBWV (but without specifying either of the currently recognized species BBWV-1 or BBWV-2) was reported affecting plants of Exacum affine in Japan (Natsuaki et al. 2001). The virus-infected Exacum plants show severe dwarfing, uprolling of the leaves, and both reduced number and reduced size of flowers. The virus was shown to be transmitted non-persistently by Myzus persicae and is also transmissible by mechanical sap-inoculation. For more details of BBWV, refer to Vicia faba.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

E Family: Tospoviridae

INSV infection in plants of Exacum spp. was reported from the USA and Japan (Daughtrey 1996; Daughtrey et al. 1997; Goto et al. 2001). The virus-infected Exacum plants exhibit symptoms of stem lesions/chlorotic ringspots, distortion of young growth, stunting, and wilting. These symptoms can vary with the stage of growth of the host and with a variety of cultural conditions. The virus is transmitted by a thrips vector Frankliniella occidentalis in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Exacum spp. was reported from the USA (Jones and Moyer 1987; Daughtrey 1996). The virus-infected Exacum plants show straw-colored necrotic leaf lesions of 7–14 mm diameter. Leaves may become completely necrotic and collapse. Stem lesions become slightly sunken and light to dark gray or straw colored. On larger plants, one branch or the entire plant may collapse and die. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Daughtrey ML (1996) Detection and identification of tospoviruses in greenhouses. Acta Hortic 431:90–98 Daughtrey M, Jones RK, Moyer JW, Daub ME, Baker JR (1997) Tospoviruses strike the greenhouse industry – INSV has become a major pathogen on flower crops. Plant Dis 81:1220–1230 Goto T, Sazarashi H, Nozawa H, Nakayama K, Natsuaki T (2001) Necrotic spot disease of cyclamen (Cyclamen persicum) and exacum (Exacum affine) caused by Impatiens necrotic spot virus (INSV) (in Japanese with english summary). Ann Rep Kanto-Tosan Plant Prot Soc 48:97–100 Jones RK, Moyer JW (1987) Exacum, a new host for tomato spotted wilt virus. NC Flower Grow Bull 31:1–2 Natsuaki KT, Kurihara J, Tomaru K (2001) Exacum (Gentianaceae) dwarf, a new disease caused by broad bean wilt virus. J. Agric. Sci: Tokyo Univ Agric 46:79–83

998

Exomis microphylla

Exomis microphylla Family: Amaranthaceae

Ornamental

Exomis microphylla latent virus Taxonomic position Genus: Becurtovirus

(EmLV)

Family: Geminiviridae

Geographical distribution EmLV infection in plants of Exomis microphylla was reported from South Africa (Claverie et al. 2018). Transmission The virus is transmitted by leahopper vectors in a persistent manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is a single molecule of circular single stranded DNA of 2974 nt (MG001960 = NC_037065) (Claverie et al. 2018; Zerbini et al 2018).

References Claverie S, Bernardo P, Kraberger S, Hartnady P, Lefeuvre P, Lett JM, Galzi S, Filloux D, Harkins GW, Varsani A, Martin DP, Roumagnac P (2018) From spatial metagenomics to molecular characterization of plant viruses: a geminivirus case study. Adv Virus Res 101:55–83 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: geminiviridae. J Gen Virol 98:131–133

F

Fagus sylvatica (European beech) Family: Fagaceae

Trees/Shrubs

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Fagus sylvatica was reported from Germany (Winter and Nienhaus 1989). The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Brome mosaic virus Taxonomic position Genus: Bromovirus

(BMV)

Family: Bromoviridae

BMV infection in plants of Fagus sylvatica was reported from Germany (Winter and Nienhaus 1989; Hamacher and Quadt 1994). The virus is transmitted by beetle vectors and is also transmissible by mechanical sap-inoculation. The virus is seed-transmitted in Fagus sylvatica plants up to 24% (Hamacher and Quadt 1994). For more details of BMV, refer to Bromus spp.

Cherry leaf roll virus Taxonomic position Genus: Nepovirus

(CLRV)

Family: Secoviridae

CLRV infection in plants of Fagus sylvatica was reported from Germany (Jones et al. 1990; Hamacher and Quadt 1994; Werner et al. 1997; von Bargen et al. 2009). The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation, and by © Springer Nature India Private Limited 2019 K. S. Sastry et al., Encyclopedia of Plant Viruses and Viroids, https://doi.org/10.1007/978-81-322-3912-3

Farfugium japonicum (Leopard plant)

1000

grafting. The virus is seed-transmitted in Fagus sylvatica plants up to 17% (Winter and Nienhaus 1989; Hamacher and Quadt 1994). For more details of CLRV, refer to Prunus avium.

Potato virus X

(PVX)

Taxonomic position Genus: Potexvirus

Family: Alphaflexiviridae

PVX infection in plants of Fagus sylvatica was reported from Germany (Winter and Nienhaus 1989). The virus is transmissible by mechanical sap-inoculation, and also by contact between plants. For more details of PVX, refer to Solanum tuberosum.

References Hamacher J, Quadt A (1994) Isolation of cherry leaf roll and brome mosaic viruses from European beech and transmission to beech seedlings. Plant Dis 78:849–853 Jones AT, Koenig R, Lesemann D-E, Hamacher J, Nienhaus F, Winter S (1990) Serological comparison of isolates of cherry leaf roll virus from diseased beech and birch trees in a forest decline area in Germany with other isolates of the virus. J Phytopathol 129:339–344 von Bargen S, Grubits E, Jalkanen R, Büttner C (2009) Cherry leaf roll virus – an emerging virus in Finland? Silva Fenn 43:727–738 Werner R, Mühlbach H-P, Büttner C (1997) Detection of cherry leaf roll nepovirus (CLRV) in birch, beech and petunia by immunocapture RT-PCR using a conserved primer pair. Eur J For Pathol 27:309–318 Winter S, Nienhaus F (1989) Identification of viruses from European beech (Fagus sylvatica L.) of declining forests in Northrhine-Westfalia (FRG). Eur J For Pathol 19:111–118

Farfugium japonicum (Leopard plant) Family: Asteraceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Farfugium japonicum was reported from Korea (Yoon et al. 2017). The virus-infected leopard plants exhibit symptoms of mosaic with dark green islands surrounding the veins. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

References Yoon JY, Choi GS, Choi SK (2017) First report of Cucumber mosaic virus in Farfugium japonicum in Korea. Plant Dis 101:264

Festuca spp.

1001

Felicia amelloides (Blue daisy) Family: Asteraceae

Ornamental

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Felicia amelloides was reported from Japan (Uga and Tsuda 2005). The virus-infected blue daisy plants exhibit chlorotic rings and necrotic spot symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Uga H, Tsuda S (2005) A one-step reverse transcription-polymerase chain reaction system for the simultaneous detection and identification of multiple tospovirus infections. Phytopathology 95:166–171

Festuca spp. Family: Poaceae

Forage crop

Barley yellow dwarf virus PAV Taxonomic position Genus: Luteovirus

(BYDV-PAV)

Family: Luteoviridae

BYDV-PAV infection in plants of Festuca spp. was reported from the USA, France, Tasmania, and South Australia (Weerapat et al. 1972; Sechler 1982; Grafton et al. 1982; Guy et al. 1986; Henry and Dedryver 1991; Henry et al. 1992). The virus-infected festuca plants do not exhibit any symptoms. The virus is transmitted by aphid vectors including Rhopalosiphon padi and Sitobion avenae in a circulative, non-propagative manner (Sechler 1982). The virus is not transmissible by mechanical sapinoculation. For more details of BYDV-PAV, refer to Hordeum vulgare.

Barley yellow dwarf virus MAV Taxonomic position Genus: Luteovirus

(BYDV-MAV)

Family: Luteoviridae

BYDV-MAV infection in plants of Festuca novae-zelandiae was reported from New Zealand (Delmiglio et al. 2010). The virus is transmitted by a aphid vectors in a circulative non-propagative

F

Festuca spp.

1002

manner. The virus is not transmitted by mechanical sap-inoculation. For more details of BYDV-MAV, refer to Hordeum vulgare.

Brome mosaic virus Taxonomic position Genus: Bromovirus

(BMV)

Family: Bromoviridae

BMV-infected plants of Festuca spp. was reported from Lithuania (Urbanaviciene and Zizyte 2012). The virus-infected festuca plants exhibit mosaic mottling and streaks on leaves and stems. The virus is transmitted by beetle vectors and is also transmissible by mechanical sap-inoculation. For more details of BMV, refer to Bromus spp.

Cereal yellow dwarf virus RPV Taxonomic position Genus: Polerovirus

(CYDV-RPV)

Family: Luteoviridae

CYDV-RPV infection in plants of Festuca spp. was reported from the USA and South Australia (Weerapat et al. 1972; Grafton et al. 1982; Henry et al. 1992). The virus-infected festuca plants do not exhibit any symptoms. The virus is transmitted by the aphid vectors including Rhopalosiphum padi, in a circulative, non-propagative manner (Grafton et al. 1982). The virus is not transmitted by mechanical sap-inoculation. For more details of CYDV-RPV, refer to Hordeum vulgare.

Festuca leaf streak cytorhabdovirus Taxonomic position Genus: Cytorhabdovirus

(FLSV)

Family: Rhabdoviridae

Geographical distribution FLSV infection in plants of Festuca spp. was reported from Denmark and Germany (Lundsgaard and Albrechtsen 1976; Lundsgaard 1984). Symptoms and host(s) The virus-infected festuca plants exhibit symptoms of bright green, elongated streaks along the veins on the leaves, mosaic, and mild leaf streaking. Festuca gigantea is the only known natural host of this virus. Transmission The virus is transmitted by the planthopper vector Javesella pellucida in a persistent-propagative manner (Lundsgaard 1999). The virus is not mechanically sap-transmissible nor is it transmitted through seed. Virion properties and genome The virions are enveloped, bacilliform, and measure 61 nm in diameter and 330 nm long. There are distinct surface projections (spikes) dispersed evenly over all the surface. The genome is a monopartite,

Festuca spp.

1003

negative-sense, single-stranded RNA (Lundsgaard and Albrechtsen 1976, 1979; Dietzgen 2011; Walker et al. 2018).

Festuca necrosis virus

(FNV)

Taxonomic position FNV is a tentative member of the genus Closterovirus and family Closteroviridae Geographical distribution FNV infection in plants of Festuca pratensis was reported from Germany and Lithuania (Schmidt et al. 1963; Urbanavieiene 2004). Symptoms and host(s) The virus-infected festuca plants exhibit symptoms of chlorosis on emerging leaves. Symptoms progress as irregularly shaped necrotic areas, combined with death of shoots or the entire plant from the roots upward. Transmission The virus is transmitted by an aphid vector Rhopalosiphum padi. Virion properties and genome The virions are flexuous filaments with an average length of 1725 nm and 18 nm in wide. The genome is presumed to be positive sense ssRNA.

Ryegrass mosaic virus Taxonomic position Genus: Rymovirus

(RGMV)

Family: Potyviridae

RGMV infection in plants of Festuca pratensis was reported from Lithuania (Urbanaviciene 2004, 2008). The virus-infected festuca plants exhibit symptoms of mosaic spotting and chlorotic and necrotic streaks on leaves and stems. The virus is transmitted by the mite vector Abacarus hystrix in a semipersistent manner, and is also transmissible through mechanical sap-inoculation. For more details of RGMV, refer to Lolium spp.

References Delmiglio C, Pearson MN, Lister RA, Guy PL (2010) Incidence of cereal and pasture viruses in New Zealand’s native grasses. Ann Appl Biol 157:25–36 Dietzgen RG (2011) Cytorhabdovirus. Rhabdoviridae. In: The Springer index of viruses. Springer, New York, pp 1709–1713. https://doi.org/10.1007/978-0-387-95919-1_277 Grafton KF, Poehlman JM, Sehgal OP, Sechler DT (1982) Tall fescue as a natural host and aphid vectors of Barley yellow dwarf virus in Missouri. Plant Dis 66:318–320 Guy PL, Johnstone GR, Duffus JE (1986) Occurrence and identity of barley yellow dwarf viruses in Tasmanian pasture grasses. Aust J Agric Res 37:43–53 Henry M, Dedryver CA (1991) Occurrence of barley yellow dwarf virus in pastures of western France. Plant Pathol 40:93–99

F

Festulolium loliaceum

1004

Henry M, Francki RIB, Wallwork H (1992) Occurrence of barley yellow dwarf virus in cereals and grasses of the lowrainfall wheat belt of South Australia. Plant Pathol 41(6):713–721 Lundsgaard T (1984) Comparison of Festuca leaf streak virus antigens with those of three other rhabdoviruses infecting the Gramineae. Intervirology 22:50–55 Lundsgaard T (1999) Javesella pellucida (F.) is a vector of Festuca leaf streak virus (FLSV, genus Cytorhabdovirus). J Plant Dis Protect 106:545–549 Lundsgaard T, Albrechtsen SE (1976) Electron microscopy of rhabdovirus-like particles in Festuca gigantea with leaf mosaic. Phytopathol Z 87:12–76 Lundsgaard T, Albrechtsen SE (1979) Ultrastructure of Festuca gigantea with rhabdovirus-like particles. Phytopathol Z 87:12–76 Schmidt HB, Richter J, Hertzsch W, Klinkowski M (1963) Untersuchungen uber eine virusbedingte Nekrose an Futtergrasern. Phytopathol Z 47:66 Sechler DT (1982) Tall fescue Festuca arundinacea as a natural host and aphid vectors of barley yellow dwarf virus in Missouri, USA. Plant Dis:318–320 Urbanaviciene L (2004) Viral diseases of poaceae family plants. Pemes Ukio Mokslai 1:19–23 Urbanaviciene L (2008) Identification of Ryegrass mosaic rymovirus in poaceae plants. Biologia 54:75–78 Urbanaviciene L, Zizyte M (2012) Identification of Brome mosaic virus in cocksfoot (Dactylis glomerata L.) and meadow fescue (Festuca pratensis Huds.) in Lithuania. Zemdirbyste-Agriculture 99:167–172 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Rhabdoviridae. J Gen Virol 99:447–448 Weerapat P, Sechler DT, Poehlman JM (1972) Host study and symptom expression of Barley yellow dwarf virus in tall fescue (Festuca arundinacea). Plant Dis Reptr 56:167–168

Festulolium loliaceum Family: Poaceae

Ryegrass mosaic virus Taxonomic position Genus: Rymovirus

Forage crop

(RGMV)

Family: Potyviridae

RGMV infection in plants of Festulolium loliaceum was reported from Lithuania (Urbanaviciene 2004, 2008). The virus-infected festulolium plants exhibit symptoms of mosaic spotting and chlorotic and necrotic streaks on leaves and stems. The virus is transmitted by the mite vector Abacarus hystrix in a semi-persistent manner, and is also transmissible through mechanical sap-inoculation. For more details of RGMV, refer to Lolium spp.

References Urbanaviciene L (2004) Viral diseases of poaceae family plants. Pemes Ukio Mokslai 1:19–23 Urbanaviciene L (2008) Identification of Ryegrass mosaic rymovirus in poaceae plants. Biologia 54:75–78

Ficus carica (Common fig)

1005

Fevillea spp. (Fevillea cordifolia and F. trilobata) Family: Cucurbitaceae

(Biodiesel) Commercial crop

Papaya ringspot virus Taxonomic position Genus: Potyvirus

(PRSV)

Family: Potyviridae

PRSV infection in plants of Fevillea spp. was reported from South America (de Castro et al. 2016). The virus-infected fevillea plants exhibit symptoms of foliar mosaic with blister-like patches. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of PRSV, refer to Carica papaya.

Zucchini yellow mosaic virus Taxonomic position Genus: Potyvirus

(ZYMV)

Family: Potyviridae

ZYMV infection in plants of Fevillea trilobata was reported from Brazil (Boiteux et al. 2013). The virus-infected fevillea plants exhibit mosaic and severe leaf malformation symptoms. Apical mosaic was slightly more severe in female than in male plants. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ZYMV, refer to Cucurbita pepo.

References Boiteux LS, Spadotti DMA, Rezende JAM, Kitajima EW (2013) Fevillea trilobata as a natural host of Zucchini yellow mosaic virus in Brazil. Plant Dis 97:1261 de Castro GM, Lima MF, Fonseca MEN, Boiteux LS, Lobo FP, Silva FR, Rech E (2016) First report of Papaya ringspot virus – type W infecting Fevillea species (Cucurbitaceae) in South America. Plant Dis 100:2540

Ficus carica (Common fig) Family: Moraceae

Edible fruit

Apple dimple fruit viroid Taxonomic position Genus: Apscaviroid

(ADFVd)

Family: Pospiviroidae

F

Ficus carica (Common fig)

1006

ADFVd was detected in plants of Ficus carica from Italy (Chiumenti et al. 2014). The viroid-infected common fig plants exhibit severe leaf malformation and mosaic symptoms. The viroid is transmissible by mechanical sap-inoculation, and also by grafting. Viroid-infected planting material is one of the main sources of this viroid spread. For more details of ADFVd, refer to Malus domestica.

Citrus exocortis viroid Taxonomic position Genus: Pospiviroid

(CEVd)

Family: Pospiviroidae

CEVd infection in plants of Ficus carica was reported from Tunisia and Iran (Yakoubi et al. 2007; Amiri Mazhar et al. 2012). The viroid-infected common fig plants exhibit mosaic symptoms. The viroid is mechanically sap-transmissible including through the use of contaminated tools. The use of infected budwood and root stocks is the primary mode of spread. For more details of CEVd, refer to Citrus spp.

Fig badnavirus 1 Taxonomic position Genus: Badnavirus

(FBV-1)

Family: Caulimoviridae

Geographical distribution FBV-1 is known to occur in Ficus carica trees in different countries worldwide including Europe, South and North America, New Zealand, Australia, Croatia, and Iran (Tzanetakis et al. 2010; Elci et al. 2012; Laney et al. 2012; Minafra et al. 2012a, b; Alimoradian et al. 2014; Alkowni et al. 2015; Voncina et al. 2015; Delic et al. 2017).

Symptoms and host(s) The virus-infected common fig plants exhibit symptoms of mottling and chlorotic spots with necrotic margins on the leaves.

Transmission The virus is mechanically sap-transmissible to seven other herbaceous plant species from different families.

Virion properties and genome The virions are bacilliform in shape with a diameter of 30 nm and modal particle length of 130 nm. The genome consists of a circular, double-stranded DNA molecule of 7140 bp (NC_017830, JF411989) with three open reading frames (ORF). ORFIII encodes a large polyprotein containing movement protein, capsid protein, aspartic protease, reverse transcriptase (RT), and RNaseH (RH) (Laney et al. 2012; Olszewski and Lockhart 2011; Bhat et al. 2016). The viral genome is integrated into the fig genome in at least some fig accessions (Laney et al. 2012).

Ficus carica (Common fig)

Fig cryptic virus

1007

(FCV)

Taxonomic position Genus: Deltapartitivirus

Family: Partitiviridae

Geographical distribution FCV infection in plants of Ficus carica was reported from Albania, Algeria, Italy, Lebanon, Syria, Turkey, and Tunisia (Elbeaino et al. 2011a; Elci et al. 2012; El Air et al. 2015). Symptoms and host(s) The virus-infected common fig plants do not exhibit any symptoms. Transmission The virus is not mechanically sap-transmissible. The virus is not transmissible by grafting. Virion properties and genome The virions are isometric, non-enveloped, and about 40 nm in diameter. The genome consists of two molecules of linear dsRNA: RNA1 consists of 1696 bp (FR687854 = NC_015494) and RNA2 of 1415 bp (FR687855 = NC_015495). Each genomic component encodes a single protein: a 54 kDa RNA-dependent RNA polymerase by RNA 1 and a 38 kDa, viral coat protein by RNA 2 (Elbeaino et al. 2011a; Ale-Agha and Rakhshandehroo 2014; Vainio et al. 2018).

Fig fleck-associated virus

(FFkaV)

Taxonomic position FFkaV is a tentative member of the genus Maculavirus and family Tymoviridae. Geographical distribution FFkaV infection in plants of Ficus carica was reported from Tunisia, China, Iran, and Syria (Elbeaino et al. 2011b, 2012b; Ale-Agha et al. 2013, 2014; El Air et al. 2015; Delic et al. 2017). Symptoms and host(s) The virus-infected common fig plants exhibit mosaic symptoms. Transmission There is no known vector for this virus. Virion properties and genome The genome is a positive-sense, single-stranded RNA of 6704 nt (NC_015229) excluding the 30 -terminal poly(A) tract and contains two open reading frames (Elbeaino et al. 2011b; Ale-Agha et al. 2014; He et al. 2017).

Fig latent virus 1

(FLV-1)

Taxonomic position FLV-1 is a tentative member of the genus Trichovirus and family Betaflexiviridae

F

Ficus carica (Common fig)

1008

Geographical distribution FLV-1 infection in plants of Ficus carica was reported from Italy, Turkey, Greece, Bosnia-Herzegovina, Montenegro, Hungary, Portugal, France, England, California, Mexico, Tunisia, Syria, Saudi Arabia and South Africa (Elci et al. 2012; Elbeaino et al. 2012b; El Air et al. 2013, 2015; Alkowni et al. 2015; Elbeshehy et al. 2017). Symptoms and host(s) The virus-infected common fig trees were symptomless or showed mosaic symptoms (Gattoni et al. 2009). Transmission The virus has no known vector. The virus is seed-transmitted at low frequency (Mink 1993; Johansen et al. 1994; Castellano et al. 2009). The virus is transmissible by mechanical sap-inoculation to a very restricted range of herbaceous hosts without inducing apparent symptoms. Use of infected vegetative planting material is the primary source of virus spread. Virion properties and genome The virions are very flexuous filaments, 700 nm long, and 10–12 nm in size. The genome consists of a single molecule of a linear, positive-sense, ssRNA about 7.5–8.0 kb in size (Gattoni et al. 2009). A partial sequence (6620 nt) lacking the 50 and 30 regions is available (FN377573) (Elbeshehy et al. 2017).

Fig leaf mottle-associated virus 1

(FLMaV-1)

Taxonomic position FLMaV-1 is a tentative member of the genus Closterovirus and family Closteroviridae Geographical distribution FLMaV-1 infection in plants of Ficus carica was first identified in the province of Taranto (Apulia, southern Italy) (Elbeaino et al. 2006). The virus spreads in Tunisia, Lebanon, Egypt, Saudi Arabia, Mexico, South Africa, New Zealand, and Turkey (Nahdi et al. 2006; Castellano et al. 2007; Elbeaino et al. 2007a; Minafra et al. 2009, 2012b; Caglar et al. 2011; Elbeshehy and Elbeaino 2011; Elbeaino et al. 2011c; Fayez and Mahmoud 2011; Alhudaib 2012; Elbeaino et al. 2012c; Elci et al. 2012; AlfaroFernandez et al. 2014; Aldhebiani et al. 2015; El Air et al. 2015; Delic et al. 2016, 2017). Symptoms and host(s) The virus-infected common fig plants exhibit symptoms of chlorotic mottling and vein-clearing of the leaves. Transmission The virus is not sap-transmissible to herbaceous hosts. Virion properties and genome The virions are filaments virus particles of up to 1800 nm and were readily seen in symptomatic leaves. Virions were observed in phloem parenchyma and companion cells but not in mesophyll cells. The genome consists of a single molecule of positive-sense ssRNA. Partial gene sequence of the heat shock

Ficus carica (Common fig)

1009

protein 70 homologue containing 616 nt (AM113547) provided evidence that FLMaV-1 is a member of the family Closteroviridae (Elbeaino et al. 2006, 2009a).

Fig leaf mottle-associated virus 2

(FLMaV-2)

Taxonomic position FLMaV-2 is a tentative member of the genus Ampelovirus and family Closteroviridae Geographical distribution FLMaV-2 infection in plants of Ficus carica was reported from Iran, Algeria, Lebanon, Tunisia, Saudi Arabia, and Turkey (Elbeaino et al. 2007a, b, 2009a; Minafra et al. 2009; Caglar et al. 2011; Elbeshehy and Elbeaino 2011; Elbeaino et al. 2012b; Elbeaino et al. 2012c; Elci et al. 2012; Danesh-Amuz et al. 2013; Aldhebiani et al. 2015; Alkowni et al. 2015; El Air et al. 2015; Delic et al. 2017). Symptoms and host(s) The virus-infected common fig plants exhibit symptoms of chlorotic mottling and vein-clearing of the leaves. Transmission The virus is not sap-transmissible to herbaceous hosts. Virion properties and genome The virions are filaments and about 2100  12 nm in size. The genome consists of a single molecule of positive-sense ssRNA; a partial genome sequence of 5868 nt (FJ473383) is available. Partial gene sequence of the heat shock protein 70 homologue showed that FLMaV-2 is related to members of the genus Ampelovirus (Elbeaino et al. 2007b).

Fig mild mottle-associated virus

(FMMaV)

Taxonomic position FMMaV is a tentative member of the genus Closterovirus and family Closteroviridae. Geographical distribution FMMaV infection in plants of Ficus carica was reported from Italy, Tunisia, Syria, New Zealand, Saudi Arabia, Lebanon, and Turkey (Elbeaino et al. 2010, 2012b; Elbeshehy and Elbeaino 2011; Elbeaino et al. 2012c; Elci et al. 2012; Minafra et al. 2012b; El Air et al. 2013, 2015; Alfaro-Fernandez et al. 2014; Aldhebiani et al. 2015; Delic et al. 2016, 2017). Symptoms and host(s) The virus-infected common fig plants exhibit symptoms of light mottling and mild malformation of the leaves. Transmission The virus was not transmissible to herbaceous hosts by mechanical inoculation.

F

Ficus carica (Common fig)

1010

Virion properties and genome The virions have filaments morphology and measure up to ca. 2000 nm. The genome consists of a single molecule of positive-sense ssRNA. A partial genome sequence of 6290 nt (FJ611959) covering seven open reading frames revealed it as a member of genus Closterovirus (Elbeaino et al. 2010).

Fig mosaic emaravirus Taxonomic position Genus: Emaravirus

(FMV)

Family: Fimoviridae

Geographical distribution Fig mosaic disease a long-known malady of fig (Ficus carica) was first reported from California (Condit and Horne 1933). FMV occurs wherever the fig crop is grown, including North America, Australia, Asia, Africa, and Europe; including China, Egypt, Turkey, Iran, Italy, Mexico, New Zealand, Tunisia, Saudi Arabia, Syria, Spain, Lebanon, and Croatia (Nitta et al. 1995; Caglar et al. 2011; Shahmirzaie et al. 2010; Elbeshehy and Elbeaino 2011; Elbeaino et al. 2011c; Alhudaib 2012; Elci et al. 2012; El Air et al. 2012, 2015; Elbeaino et al. 2012a; Elbeaino et al. 2012c; Ishikawa et al. 2012a; Minafra et al. 2012b; AlfaroFernandez et al. 2014; Mijit et al. 2015; Aldhebiani et al. 2015; Alkowni et al. 2015; Voncina et al. 2015; Delic et al. 2016, 2017). Symptoms and host(s) The virus-infected common fig plants exhibit symptoms on leaves of mosaic and ringspots which are distinctly yellow, contrasting with normal green color of the foliage. The margins of the yellow spots blend gradually from a light yellow color into the dark green of healthy tissue. Mosaic spots or lesions may be uniformly scattered over the surface of the leaves or may appear as irregular patches of light green diffused widely throughout the leaf blade. Later in the season, a rust-colored band develops along the border of the mosaic spots, apparently caused by the death of epidermal or subepidermal cells. Deformed leaves may occur on the same twig as normal leaves. Yellow spots on fruits are very conspicuous. Premature fruit drop may also occur in certain cultivars. The natural host range is apparently restricted to Ficus carica (Omcikus and Blagojevic 1957). Transmission The virus is transmitted by the eriophyid mite, Aceria ficus (Flock and Wallace 1955; Caglayan et al. 2012). The virus spreads naturally with infected cuttings and suckers by which fig is exclusively propagated. Virion properties and genome The virions are quasi-spherical, double-membrane-bound particles (DMBs) of 80–100 nm. The genome consists of six negative sense ssRNA segments: RNA1 consists of 7039 nt (AM941711 = NC_029562), RNA2 of 2252 nt (FM864225 = NC_029565), RNA3 of 1490 nt (FM991954 = NC_029563), RNA4 of 1472 nt (FM992851 = NC_029564), RNA5 of 1752 nt (HE803826 = NC_029566), and RNA6 of 1212 nt (HE803827 = NC_029568) (Elbeaino et al. 2009b, c, 2012a; Ishikawa et al. 2012b, 2013; Mielke-Ehret and Muhlbach 2012; Elbeaino et al. 2018).

Ficus carica (Common fig)

1011

Hop stunt viroid

(HpSVd)

Taxonomic position Genus: Hostuviroid

Family: Pospiviroidae

HpSVd was detected from plants of Ficus carica in Syria and Iran (Yakoubi et al. 2007; Amiri Mazhar et al. 2012; Elbeaino et al. 2012b, 2013). The viroid is mechanically sap-transmissible. The use of shoots from the infected plants for propagation is the primary cause of spread of this viroid. For more details of HpSVd, refer to Humulus lupulus.

F References Aldhebiani AY, Elbeshehy EKF, Baeshen AA, Elbeaino T (2015) Four viruses infecting figs in Western Saudi Arabia. Phytopathol Mediterr 54:497–503 Ale-Agha GN, Rakhshandehroo F (2014) Detection and molecular variability of Fig fleck-associated virus and Fig cryptic virus in Iran. J Phytopathol 162:417–425 Ale-Agha GN, Rakhshandehroo F, Zamanizadeh HR, Elbeaino T (2013) Presence of fig fleck-associated virus in fig trees in Iran. J Plant Pathol 95:664 Alfaro-Fernandez A, Hernández-Llopis D, Font MI (2014) Fig viruses in Mainland Spain. J Phytopathol 162:332–337 Alhudaib K (2012) Incidence of fig mottle-associated virus and fig mosaic virus in Eastern province of Saudi Arabia. Int J Virol 8:128–132 Alimoradian MR, Rakhshandehroo F, Shams-bakhsh M (2014) First record of Fig badnavirus-1 in fig trees in Iran. J Plant Pathol 96:S4.113–S4.131 Alkowni R, Chiumenti M, Minafra A, Martelli GP (2015) A survey for fig-infecting viruses in Palestine. J Plant Pathol 97(2):383 Amiri Mazhar M, Bagherian SAA, Izadpanah K (2012). Identification and isolation of hop stunt viroid and citrus exocortis viroid from fig in Iran. Paper presented at the 20th Iranian Plant Protection Congress, Shiraz, Iran, 25–28 August Bhat AI, Hohn T, Selvarajan R (2016) Badnaviruses: the current global scenario. Viruses 8(6):177 Caglar BK, Fidan H, Güldür ME, Elbeaino T (2011) The prevalence of three viruses infecting fig in southern Turkey. J Phytopathol 159:181–183 Caglayan K, Elci K, Ulubas Serce C, Kaya K, Gazel M, Medina V (2012) Detection of Fig mosaic virus in viruliferous eriophyid mite Aceria ficus. J Plant Pathol 94:629–634 Castellano MA, Gattoni G, Minafra A, Conti M, Martelli GP (2007) Fig mosaic in Mexico and South Africa. J Plant Pathol 89:441–444 Castellano MA, De Stradis A, Minafra A, Boscia D, Martelli GP (2009) Seed transmission of Fig latent virus 1. J Plant Pathol 91:697–700 Chiumenti M, Torchetti EM, Di Serio F, Minafra A (2014) Identification and characterization of a viroid resembling Apple dimple fruit viroid in fig (Ficus carica L.) by next generation sequencing of small RNAs. Virus Res 188:54–59 Condit IJ, Horne WT (1933) A mosaic of the fig in California. Phytopathology 23:887–896 Danesh-Amuz S, Rakhshandehroo F, Rezaee S, Elbeaino T (2013) Occurrence of fig leaf mottle-associated virus 2 in Iran. J Plant Pathol 95:663 Delic D, Elbeaino T, Lolic B, Duric G (2016) Detection of Fig mosaic virus, Fig leaf mottle-associated virus 1, and Fig mild mottle-associated virus in Bosnia and Herzegovina. J Plant Pathol 98:371 Delic D, Perovic T, Hrncic S, Lolic B, Duric G, Elbeaino T (2017) Detection and phylogenetic analyses of fig-infecting viruses in Bosnia and Herzegovina and Montenegro. Phytopathol Mediterr 56:470–478 El Air M, Mahfoudhi N, Elbeaino T, Dhouibi MH, Digiaro M (2012) Occurrence of Fig mosaic virus (FMV) in Tunisian fig orchards. J Plant Pathol 94:S4.89 El Air M, Mahfoudhi N, Elbeaino T, Dhouibi MH, Digiaro M (2013) Presence of fig mild mottle-associated virus and fig latent virus-1 in Tunisia. J Plant Pathol 95:S4.69–S4.77 El Air M, Mahfoudhi N, Digiaro M, Dhouibi MH, Elbeaino T (2015) Incidence and distribution of viruses in Tunisian fig orchards. J Plant Pathol 97(2):327–331. https://doi.org/10.4454/JPP.V97I2.030 Elbeaino T, Digiaro M, De Stradis A, Martelli GP (2006) Partial characterization of a closterovirus associated with a chlorotic mottling of fig. J Plant Pathol 88:187–192

1012

Ficus carica (Common fig)

Elbeaino T, Choueiri E, Hobeika C, Digiaro M (2007a) Presence of Fig leaf mottle associated virus 1 and 2 in Lebanese fig orchards. J Plant Pathol 89:409–411 Elbeaino T, Digiaro M, De Stradis A, Martelli GP (2007b) Identification of a second member of the family Closteroviridae in mosaic-diseased figs. J Plant Pathol 89:119–124 Elbeaino T, Nahdi S, Digiaro M, Alabdullah A, Martelli GP (2009a) Detection of Fig leaf mottle-associated virus 1 and Fig leaf mottle-associated virus 2 in the Mediterranean region and study on sequence variation of the hsp70 gene. J Plant Pathol 91:425–431 Elbeaino T, Digiaro M, Martelli GP (2009b) Complete nucleotide sequence of four RNA segments of Fig mosaic virus. Arch Virol 154:1719–1727 Elbeaino T, Digiaro M, Alabdullah A, De Stradis A, Minafra A, Mielke N, Castellano MA, Martelli GP (2009c) A multipartite single-stranded negative-sense RNA virus is the putative agent of fig mosaic disease. J Gen Virol 90(PT 5):1281–1288 Elbeaino T, Digiaro M, Heinoun K, De Stradis A, Martelli GP (2010) Fig mild mottle-associated virus, a novel closterovirus infecting fig. J Plant Pathol 92:165–172 Elbeaino T, Abou KR, Digiaro M, Minafra A, Martelli GP (2011a) The complete nucleotide sequence and genome organization of Fig cryptic virus, a novel bipartite dsRNA virus infecting fig, widely distributed in the Mediterranean basin. Virus Genes 42:415–421 Elbeaino T, Digiaro M, Martelli GP (2011b) Complete sequence of Fig fleck-associated virus, a novel member of the family Tymoviridae. Virus Res 161:198–202 Elbeaino T, Gonzalez Rodriguez AM, Grajal-Martin MJ, Digiaro M (2011c) Survey of fig viruses in the Canary Islands. J Plant Pathol 93(3):737–739 Elbeaino T, Digiaro M, Martelli GP (2012a) RNA-5 and -6, two additional negative-sense RNA segments associated with Fig mosaic virus. J Plant Pathol 94:421–425 Elbeaino T, Abou Kubaa R, Ismaeil F, Mando J, Digiaro M (2012b) Viruses and Hop stunt viroid of fig trees in Syria. J Plant Pathol 94:687–691 Elbeaino T, Mortada C, Digiaro M (2012c) Survey on fig viruses in Lebanon. Acta Hortic 940:665–668 Elbeaino T, Choueiri E, Digiaro M (2013) First report of hop stunt viroid in Lebanese fig trees. J Plant Pathol 95:218 Elbeaino T, Digiaro M, Mielke-Ehret N, Muelbach H-P, Martelli GP, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Fimoviridae. J Gen Virol 99:1478–1479 Elbeshehy EKF, Elbeaino T (2011) Viruses infecting figs in Egypt. Phytopathol Mediterr 50:327–332 Elbeshehy EKF, Adhebiani AY, Hassasn WM (2017) Partial molecular characterization of the Fig latent virus 1 (FLV-1) infecting figs in Western Saudi Arabia. Res J Biotech 12(3):91–98 Elci E, Serce C, Gazel M, Caglayan K (2012) Molecular detection and comparative sequence analysis of viruses infecting fig trees in Turkey. J Phytopathol 160:418–423 Fayez KA, Mahmoud SY (2011) Detection and partial characterization of a putative closterovirus affecting Ficus carica: molecular, ultrastructural and physiological aspects of infected leaves. Acta Physiol Plant 33:2187–2198 Flock RA, Wallace JM (1955) Transmission of fig mosaic by eriophyid mite Aceria ficus. Phytopathology 45:52–54 Gattoni G, Minafra A, Castellano MA, De Stradis A, Boscia D, Elbeaino T, Digiaro M, Martelli GP (2009) Some properties of Fig latent virus 1, a new member of the family Flexiviridae. J Plant Pathol 91:555–564 He Z, Mijit M, Li S, Zhang Z (2017) Complete nucleotide sequence of a novel strain of fig fleck-associated virus from China. Arch Virol 162:1145–1148 Ishikawa K, Maejima K, Nagashima S, Sawamura N, Takinami Y, Komatsu K, Hashimoto M, Yamaji Y, Yamamoto J, Namba S (2012a) First report of Fig mosaic virus infecting common fig (Ficus carica) in Japan. J Gen Plant Pathol 78:136–139 Ishikawa K, Maejima K, Komatsu K, Kitazawa Y, Hashimoto M, Takata D, Yamaji Y (2012b) Identification and characterization of two novel genomic RNA segments of Fig mosaic virus, RNA5 and RNA6. J Gen Virol 93:1612–1619 Ishikawa K, Maejima K, Komatsu K, Netsu O, Keima T, Shiraishi T, Okano Y, Hashimoto M, Yamaji Y, Namba S (2013) Fig mosaic emaravirus p4 protein is involved in cell-to-cell movement. J Gen Virol 94:682–686 Johansen E, Edwards MC, Hampton RO (1994) Seed transmission of viruses: current perspectives. Annu Rev Phytopathol 32:363–386 Laney AG, Hassan M, Tzanetakis IE (2012) An integrated badnavirus is prevalent in fig germplasm. Phytopathology 102:1182–1189 Mielke-Ehret N, Muhlbach HP (2012) Emaravirus: a novel genus of multipartite, negative strand RNA plant viruses. Viruses 4:1515–1536 Mijit M, Li SF, Zhang S, Zhang ZX (2015) First report of Fig mosaic virus infecting common fig (Ficus carica) in China. Plant Dis 99:422.3 Minafra A, De Stradis A, Clara MI, Martelli GP (2009) Viruses in mosaic affected figs in Portugal and the Island of Crete. J Plant Pathol 91:238

Ficus spp. (F. benjamina, F. elastica) (Weeping fig; Rubber fig)

1013

Minafra A, Chiumenti M, Elbeaino T, Digiaro M, Bottalico G, Pantaleo V, Martelli GP (2012a) Occurrence of Fig badnavirus 1 in fig trees from different countries and in symptomless seedlings. J Plant Pathol 94:S4.105 Minafra A, Chiumenti M, Martelli GP (2012b) Fig tree viruses in New Zealand. J Plant Pathol 94(4, Supplement):S4.104 Mink GI (1993) Pollen- and seed-transmitted viruses and viroids. Annu Rev Phytopathol 31:375–402 Nahdi S, Elbeaino T, Digiaro M, Martelli GP (2006) First record of Fig leaf mottle-associated virus 1 in Tunisia. J Plant Pathol 88:70 Nitta H, Imada J, Kano T, Nakamoto K, Ogasawara S (1995) Occurrence and cause of fig mosaic symptoms in Hiroshima Prefecture. Bull Hiroshima Pref Agric Res Cent 62:53–65 Olszewski NE, Lockhart B (2011) Badnavirus. Caulimoviridae. In: The Springer index of viruses. Springer, New York, pp 263–269. https://doi.org/10.1007/978-0-387-95919-1_40 Omcikus C, Blagojevic M (1957) Observations on mosaic symptoms on different varieties of fig in Herzegovina. Zasht Bilja 44:27–34 Shahmirzaie M, Rakhshandehroo F, Zamanizadeh HR, Elbeaino T, Martelli GP (2010) First report of Fig mosaic virus from fig trees in Iran. J Plant Pathol 92:S4.120 Tzanetakis IE, Laney AG, Keller KE, Martin RR (2010) New viruses found in fig exhibiting mosaic symptoms. 21st International Conference on Virus and other Graft Transmissible Diseases of Fruit Crops. Julius-Kühn-Archiv 427:79–82 Vainio EJ, Chiba S, Ghabrial SA, Maiss E, Roossinck M, Sabanadzovic S, Suzuki N, Xie J, Nibert M, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Partitiviridae. J Gen Virol 99:17–18 Voncina D, Pilipovic P, Skoric D, Krapac M, Prgomet Z, Milicevic (2015) First report of Fig mosaic virus and Fig badnavirus 1 on common fig trees in Croatia. J Plant Pathol 97:S71 Yakoubi S, Elleuch A, Besaies N, Marrakchi M, Fakhfakh H (2007) First report of hop stunt viroid and citrus exocortis viroid on fig with symptoms of fig mosaic disease. J Phytopathol 155:125–128

Ficus spp. (F. benjamina, F. elastica) (Weeping fig; Rubber fig) Family: Moraceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV was reported infecting plants of Ficus spp. in Iran (Ghotbi et al. 2005; Ghotbi and Shahraeen 2012; Ghotbi 2013). The virus-infected rubber fig plants exhibit symptoms of necrotic leaf spot, chlorosis, and stunting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV was reported infecting plants of Ficus benjamina in Iran and Spain (Lavina and Battle 1993; Ghotbi and Shahraeen 2012). The virus-infected rubber fig plants exhibit chlorotic and necrotic ringspots, leaf distortion, and tip necrosis symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large

F

Fittonia verschaffeltii (Nerve plant)

1014

number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow ring virus

(TYRV)

Synonyms Tomato Varamin virus (ToVV) Taxonomic position TYRV is a tentative member of the genus Orthotospovirus and family Tospoviridae ToVV was reported infecting plants of Ficus benjamina in Iran (Ghotbi et al. 2005). The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of TYRV, refer to Solanum lycopersicum.

References Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iran J Plant Pathol 49:41 Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses affecting propagated ornamentals in Northern Iran. Int Res J Microbiol 3:373–381 Ghotbi T, Sharaeen N, Winter S (2005) Occurrence of tospoviruses in ornamental and weed species in Markazi and Tehran provinces in Iran. Plant Dis 89:425–429 Lavina A, Battle A (1993) First report of Tomato spotted wilt virus infection of ficus species in Spain. Plant Dis 77:536

Fittonia verschaffeltii (Nerve plant) Family: Acanthaceae

Bidens mottle virus Taxonomic position Genus: Potyvirus

Ornamental

(BiMoV)

Family: Potyviridae

BiMoV was described in plants of Fittonia verschaffeltii in Florida, USA (Zettler et al. 1977; Zurawski 1979; Zurawski et al. 1980). The virus-infected nerve plants most commonly exhibit distortion of the normally symmetrical leaves. Interveinal chlorosis and stunting also occur. Symptoms on this host fluctuate seasonally, with severity greatest during the winter. The virus is transmitted by an aphid vector, Myzus persicae in a non-persistent manner, and is also transmissible by mechanical sapinoculation to species of several plant families: a number of vegetable crops are affected with this virus. For more details of BiMoV, refer to Bidens spp.

Forsythia spp.

1015

References Zettler FW, Lima JAA, Zurawski DB (1977) Bidens mottle virus infecting Fittonia spp. in Florida. Proc Am Phytopathol Soc 4:121–122 Zurawski DB (1979) Some biological and serological properties of Bidens mottle virus isolated from Fittonia. MA thesis. University of Florida, Gainesville, p 91 Zurawski DB, Purcifull DE, McRitchie JJ (1980) Bidens mottle virus of Fittonia verschaffeltii. Plant Pathology Circular No. 215. Fla. Dept. Agric. and Consumer Services

Foeniculum vulgare (Fennel) Family: Apiaceae

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

F

Medicinal

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Foeniculum vulgare was reported from Bulgaria (Dikova 2011). The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. For more details of TSWV, refer to Solanum lycopersicum.

References Dikova B (2011) Tomato spotted wilt virus on some medicinal and essential oil-bearing plants in Bulgaria. Bulg J Agric Sci 17:306–313

Forsythia spp. Family: Oleaceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

Family: Secoviridae

ArMV infection in plants of Forsythia spp. was reported from Poland (Kaminska 1985). The virusinfected forsythia plants exhibit symptoms of chlorotic patterns or yellowed leaves, leaf deformation, and stunted growth. This virus is transmitted by the nematode vectors (Xiphinema spp.) in a nonpersistent manner, and is also transmissible through mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Forsythia spp.

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Cherry leaf roll virus Taxonomic position Genus: Nepovirus

(CLRV)

Family: Secoviridae

CLRV infection in plants of Forsythia spp. was reported from Poland (Kaminska 1985). The virusinfected forsythia plants exhibit symptoms of chlorotic patterns or yellowed leaves, leaf deformation, and stunted growth. The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation, and by grafting. For more details of CLRV, refer to Prunus avium.

Cucumber mosaic virus

(CMV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV infection in plants of Forsythia koreana was reported from Korea and Poland (Kaminska 1985; Lee et al. 1997). The virus-infected forsythia plants exhibit mosaic symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sapinoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Raspberry ringspot virus Taxonomic position Genus: Nepovirus

(RpRSV)

Family: Secoviridae

RpRSV infection in plants of Forsythia spp. was reported from Poland (Kaminska 1985). The virusinfected forsythia plants exhibit symptoms of chlorosis, and reduction of leaf size. The virus is transmitted by the nematode vectors (Longidorus spp.) in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of RpRSV, refer to Rubus spp.

Tobacco necrosis virus

(TNV)

Taxonomic position TNV is a tentative member of the genus Alphanecrovirus and family Tombusviridae. TNV infection in plants of Forsythia spp. was reported from Poland (Kaminska 1985). The virusinfected forsythia plants exhibit symptoms of chlorotic patterns or yellowed leaves, leaf deformation, and stunted growth. The virus is transmitted by the zoospores of the fungus Olpidium brassicae and is also transmissible by mechanical sap-inoculation. For more details of TNV, refer to Nicotiana tabacum.

Forsythia spp.

1017

Tobacco rattle virus

(TRV)

Taxonomic position Genus: Tobravirus

Family: Virgaviridae

TRV infection in plants of Forsythia spp. was reported from Poland (Kaminska 1985). The virusinfected forsythia plants exhibit symptoms of chlorotic ringspots and patterns and sometimes reductions of the leaf blade. The virus is transmitted by nematode vectors and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

F Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Forsythia spp. was reported from Poland and the USA (Tiangco and Varney 1970; Waterworth and Povish 1972; Kaminska 1985). The virus-infected forsythia plants exhibit symptoms of chlorotic spots and rings. The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato black ring virus Taxonomic position Genus: Nepovirus

(TBRV)

Family: Secoviridae

TBRV infection in plants of Forsythia spp. was reported from Poland (Kaminska 1985). The virusinfected forsythia plants exhibit symptoms of chlorotic patterns or yellowed leaves, leaf deformation, and stunted growth. The virus is transmitted by the nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of TBRV, refer to Solanum lycopersicum.

References Kaminska M (1985) Some aspects of virus infection in Forsythia spp. In: Virus diseases of ornamentals. Acta Hortic 164:263–271 Lee SY, Park SJ, Choi JK (1997) Characterization of an isolate of cucumber mosaic virus from forsythia (Forsythia koreana Nakai.). Korean J Plant Pathol 13(5):358–363 Tiangco ES, Varney EH (1970) A vein chlorosis or yellow-net disease of Forsythia caused by Tobacco ringspot virus. Phytopathology 60:579 Waterworth HE, Povish WR (1972) Tobacco ringspot virus from naturally infected dogwood, autumn crocus and forsythia. Plant Dis Reptr 56:336–337

Fragaria spp. (Strawberry)

1018

Fragaria spp. (Strawberry) Family: Rosaceae

Edible fruit

Apple mosaic virus

(ApMV)

Taxonomic position Genus: Ilarvirus

Family: Bromoviridae

ApMV infection in plants of Fragaria spp. was reported from North America, South America, Europe, Africa, Asia, and New Zealand (Cardoni et al. 2002; Tzanetakis and Martin 2005a, 2013, 2014; Martin and Tzanetakis 2006). Newer leaves of virus-infected strawberry had a chlorotic peacock pattern with a few leaves displaying downward leaf rolling. No insect vector is known for this virus. The virus is transmissible by grafting and through the use of infected, vegetative planting material. The virus is mechanically sap-transmissible. For more details of ApMV, refer to Malus domestica.

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

Geographical distribution ArMV infection in plants of Fragaria spp. was reported from European countries (England, Scotland, Ireland, Germany), North America, South America, Africa, Asia, and New Zealand (Jha 1961; Murant 1970; Martin and Tzanetakis 2006). Symptoms and host(s) With ArMV infection of strawberry, symptoms include stunting, chlorotic leaf mottling, or mosaic; yellow spots, blotches, or streaks; and twisting, cupping, or crinkling of leaves. However, many infections remain symptomless. The virus occurs naturally in many species of wild and cultivated monocotyledonous and dicotyledonous plants. It infected 93 species in 28 dicotyledonous families when transmissible by mechanical inoculation (Schmelzer 1962), and it is also reported to infect the roots of the gymnosperm Chamaecyparis lawsoniana. It infects almost all commonly used herbaceous test plants, but isolates of the virus differ in virulence. Chenopodium amaranticolor and C. quinoa, cucumber, Nicotiana tabacum cv. White Burley, Phaseolus vulgaris cv. The Prince (French bean), and Petunia hybrida are good diagnostic hosts, which producing chlorotic or necrotic local lesions. Transmission The virus is transmitted by the nematode vectors, Xiphinema diversicaudatum and X. coxi, in a nonpersistent manner (Brown and Trudgill 1983, 1998). Once an adult nematode vector acquires the virus, it remains infective for life. However, juvenile stages do not retain the viruses through molts in the nematode life cycle, nor do females pass them on to progeny when they lay eggs. Virus spread occurs over only short distances since the nematode vectors migrate slowly through the soil. Discrete patches

Fragaria spp. (Strawberry)

1019

of infected plants are common. The virus is mechanically sap-transmissible to 93 species from 28 dicotyledonous families. Seed transmission is a common feature and was found in at least 15 species out of 12 plant families with up to nearly 100% of the progeny being infected (Murant 1970). Virion properties and genome The virions are isometric, non-enveloped of two types but similar in size, and 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is a bipartite, linear, positive-sense, single-stranded RNA. RNA1 consists of 7334 nt (AY303786 = NC_006057) and RNA2 of 3820 nt (AY017339 = NC_006056) (Loudes et al. 1995; Wetzel et al. 2001, 2004; Dunez and Le Gall 2011; Sanfacon 2015; Thompson et al. 2017).

F Beet pseudoyellows virus Taxonomic position Genus: Crinivirus

(BPYV)

Family: Closteroviridae

BPYV infection in plants of Fragaria spp. was reported from the North America including the USA, South America including Peru, Europe, Africa, Asia, and New Zealand (Tzanetakis et al. 2003a; Tzanetakis 2004; Tzanetakis and Martin 2004, 2014; Martin and Tzanetakis 2006, 2013; Wintermantel et al. 2006). The virus is often symptomless in strawberry but in some cases shows stunting and reddening of the leaves on strawberry (Fragaria  ananassa). Strawberries suffering from pallidosis disease are often mixed infections of BPYV and strawberry pallidosis-associated virus (SPaV) and older leaves develop a red color, vein and petiole reddening, roots become stunted, and plants fail to develop (Wintermantel et al. 2006). The virus is transmitted by the greenhouse whitefly (Trialeurodes vaporiorum) in a semipersistent manner. The virus is not transmissible by either mechanical sap-inoculation or contact between plants. The virus is transmissible by grafting. For more details of BPYV, refer to Beta vulgaris.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Fragaria ananassa was reported from North America, South America, Europe, Africa, Asia including China, and New Zealand (Chen et al. 2014). The virus-infected strawberry plants exhibit symptoms of leaf malformation and yellowing. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Fragaria chiloensis cryptic virus

(FCCV)

Taxonomic position FCCV is a tentative member of the genus Deltapartitivirus and family Partitiviridae

Fragaria spp. (Strawberry)

1020

Geographical distribution FCCV infection in plants of Fragaria chiloensis was reported from Chile, Mexico, and South and North America (Tzanetakis and Martin 2005b; Martin and Tzanetakis 2006; Tzanetakis et al. 2008; SilvaRosales et al. 2013). Symptoms and host(s) The virus-infected strawberry plants do not exhibit any external symptoms. Transmission The mode of transmission is unknown. Virion properties and genome The genome consists of three segments of dsRNA, with RNA 1 of 1734 bp (DQ093961 = NC_009519), RNA 2 of 1479 bp (DQ355440 = NC_009521), and RNA 3 of 1465 bp (DQ355439 = NC_009520) (Tzanetakis and Martin 2005b; Tzanetakis et al. 2008; Vainio et al. 2018).

Fragaria chiloensis latent virus Taxonomic position Genus: Ilarvirus

(FCILV)

Family: Bromoviridae

Geographical distribution FCILV infection was first reported in plants of Fragaria chiloensis from Chile by Spiegel et al. (1993). The virus spreads in North America, South America, Chile, and Mexico (Tzanetakis 2004; Martin and Tzanetakis 2006; Silva-Rosales et al. 2013; Tzanetakis and Martin 2013, 2014). Symptoms and host(s) The virus-infected strawberry plants do not exhibit any symptoms. Transmission The virus is transmissible by mechanical sap-inoculation to less than three plant families and is also transmissible through grafting. The virus is transmitted by seed up to 30–50% in Fragaria chiloensis. Virion properties and genome The virions are isometric with a modal length of 21.4 nm in diameter and up to 54.5 nm in length. The genome consists of three segments of positive-sense single-stranded RNA: RNA1 contains 3431 nt (AY682102 = NC_006566), RNA2 is 2570 nt (AY707771 = NC_006567), and RNA3 is 2484 nt (AY707772 = NC_006568) and a subgenomic RNA4 of 1.2 kb (Tzanetakis and Martin 2005c; Scott 2011a, b).

Raspberry ringspot virus Taxonomic position Genus: Nepovirus

(RpRSV)

Family: Secoviridae

Fragaria spp. (Strawberry)

1021

RpRSV-infected plants of Fragaria spp. were reported from Europe, the former USSR, Italy, and Asia (Cardoni et al. 2002; Martin and Tzanetakis 2006). The virus-infected strawberry plants show leaf chlorosis, chlorotic ringspots, rings, and line patterns. The infected plants are dwarfed and eventually die. The virus is transmitted by the nematode vector Longidorus spp. in a non-persistent manner, and there is also a report of RpRSV transmission through members of the genera Paratrichodorus and Xiphinema (Trudgill et al. 1983; Brown and Trudgill 1998). The virus is mechanically saptransmissible. The virus is seed-transmitted at a rate of between 35% and 49% in Fragaria spp. (Lister and Murant 1967). The virus is graft-transmissible, and the use of vegetative material from virus-infected plants for planting is the major mode of spread. For more details of RpRSV, refer to Rubus spp.

Strawberry chlorotic fleck-associated virus Taxonomic position Genus: Closterovirus

(SCFaV)

Family: Closteroviridae

Geographical distribution Strawberry chlorotic fleck disease was identified in 1962 by (Horn and Carver). SCFaV infection in plants of Fragaria spp. was reported from the USA (Fulton 1987b; Martin and Tzanetakis 2006; Tzanetakis and Martin 2007, 2008, 2013, 2014). Symptoms and host(s) The virus-infected commercial strawberry cultivars produce no overt symptoms. The young leaves of virus-infected Fragaria vesca are distorted and down-curled. Vein-clearing followed by the appearance of small chlorotic spots is sometimes evident. On F. virginiana plants exhibiting chlorosis, downcurling, and distortion of young leaves are observed. The virus has a narrow host range and was unable to be transmitted into any of the 11 species tested (Tzanetakis and Martin 2007). Transmission It was not possible to establish the transmission of the virus using Aphis gossypii or other species of aphids. Aphid transmission trials using the CFRA 9018 plant have been unsuccessful. This isolate has been propagated vegetatively in strawberry, possibly without any aphid transmissions for more than 40 years which may have resulted in the loss of aphid transmissibility due to accumulation of mutations (Atreya et al. 1991; Tzanetakis and Martin 2007). Experimentally the virus is transmissible through leaflet grafting to Fragaria vesca and F. virgianiana (Fulton 1987b). The virus is not mechanically sap-transmissible. Virion properties and genome The virions are flexuous filaments, non-enveloped, 700–2000 nm long, and 12 nm wide. The genome is a monopartite, linear, single-stranded, positive-sense RNA of 17,039 nt (NC_008366) and encodes ten open reading frames (Tzanetakis and Martin 2007; Agranovsky and Lesemann 2011).

Strawberry crinkle cytorhabdovirus Taxonomic position Genus: Cytorhabdovirus

(SCV)

Family: Rhabdoviridae

F

Fragaria spp. (Strawberry)

1022

Geographical distribution SCV infection in plants of F. vesca cv. Marshall was first reported from the USA (Zeller and Vaughan 1932). The virus spreads in Australia, Belgium, Bulgaria, Canada, Chile, China, the former Czechoslovakia, France, Germany, Israel, Italy, the Netherlands, New Zealand, Poland, Africa, Europe, Asia, North America, South and Central America, and the former Yugoslavia (Frazier et al. 1987; Posthuma et al. 2000; Cardoni et al. 2002; Martin and Tzanetakis 2006; Rojas et al. 2013; Perotto et al. 2014; Tzanetakis and Martin 2013, 2014). Symptoms and host(s) The virus-infected strawberry cultivars are generally symptomless. Sensitive strawberry cultivars may have dwarfed, distorted, crinkled, or otherwise deformed leaves with chlorotic to necrotic veinassociated spots, or even local lesions in petioles and stolons, with streaking and deformation of petals. Serious damage occurs in production areas where infections with both virulent and symptomless strains occur. Virulent strains severely reduce plant vigor and productivity, whereas symptomless strains (i.e., latent A) reduce plant vigor, runner formation, and fruit size and yields only in some sensitive cultivars (Freeman and Mellor 1962; McGrew and Scott 1964). The natural hosts of this virus are F. virginiana, F. chiloensis, and F. chiloensis var. ananassa. Transmission The virus is transmitted by the aphid vectors, Chaetosiphon jacobi and C. fragaefolii in a persistent, propagative manner (Posthuma et al. 2002). There is a 14-day minimum latent period from acquisition to transmission. The virus is retained when the vector moults, and multiplies in the vector; it is not transmitted congenitally to the progeny of the vector. The virus can be transmitted by injection of hemolymph from aphid and is also transmissible mechanically from strawberry to Nicotiana occidentalis and then to Physalis pubescens (Klerks et al. 2004). The virus is transmissible by grafting. The use of runners from infected strawberry plants for planting material is the major mode of spread of the virus. The virus is not transmissible by contact between plants, by seed and pollen. Virion properties and genome The virions are enveloped, bacilliform virions, 60–75 nm in diameter, and 200–350 nm long. There are distinct surface projections (spikes) dispersed evenly over all the surface. The genome is a monopartite, negative-sense, single-stranded RNA. A partial genome sequence of 1638 nt is available (AY331389) (Hunter et al. 1990; Posthuma et al. 2002; Klerks et al. 2004; Schoen et al. 2004; Dietzgen 2011). Partial sequences of the RdRp gene are available (AY250986, KJ748457); sequences of two divergent strains of SCV from the same plant have since become available (MH129615, 14,545 nt; MH129616, 14,559 nt (Koloniuk et al. 2018; Walker et al. 2018).

Strawberry latent C virus

(SLCV)

Taxonomic position SLCV is a tentative member of the genus Nucleorhabdovirus and family Rhabdoviridae. Geographical distribution SLCV infection was first reported in plants of Fragaria spp. from Japan and North America by Yoshikawa et al. (1986b). The virus spreads in the Eastern Asian region and the North American region, Canada, Japan, and the USA (McGrew 1987; Yoshikawa and Inouye 1988; EPPO/CABI 1996; Martin and Tzanetakis 2006; Tzanetakis and Martin 2013).

Fragaria spp. (Strawberry)

1023

Symptoms and host(s) The virus-infected strawberry plants are symptomless. Transmission The virus is transmitted by the aphid vector, Chaetosiphon fragaefolii, in a persistent-propagative manner. Experimentally it is also been transmissible by leaflet grafting and by dodder. Virion properties and genome The virions are rhabdo- or bullet-shaped, enveloped, of 190–380 nm in length, and 68 nm wide (from sap). The genome consists of a single-stranded, negative-sense RNA.

F Strawberry latent ringspot virus Taxonomic position Genus: Unassigned

(SLRSV)

Family: Secoviridae

Geographical distribution SLRSV infection in plants of Fragaria vesca was first reported from Scotland (Lister 1964). The virus spreads in Europe, Australia, Belgium, Canada, Finland, France, Germany, Ireland, Israel, Italy, Luxembourg, the Netherlands, New Zealand, Poland, Portugal, Romania, Spain, Switzerland, Turkey, the UK, the USA, and the former Yugoslavia (Cardoni et al. 2002; Martin et al. 2004; Martin and Tzanetakis 2006, 2013; Tzanetakis et al. 2006a). Symptoms and host(s) The virus-infected strawberry plants usually remain symptomless. Some cultivars may show mottling and decline. The virus has a wide host range that exceeds 125 plant species belonging to families representing both monocots and dicots. Transmission The virus is transmitted by the nematode vectors, Xiphinema diversicaudatum and X. coxi, (Lister 1964; Brown and Trudgill 1983, 1998). The virus is transmissible by mechanical sap-inoculation, and by grafting. It is transmissible through seeds of Rubus idaeus and number of weed hosts. Virion properties and genome The virions are isometric, 30 nm in diameter, and angular in profile (Francki et al. 1985). The genome is a bipartite, single-stranded RNA: RNA1 consists of 7496 nt (AY860978 = NC_006964) and RNA2 of 3842 nt (AY860979 = NC_006965) (Kreiah et al. 1993). The virus has two coat proteins, and the complete genome sequence has been determined (Everett et al. 1994; Tzanetakis et al. 2006a; Thompson et al. 2017).

Strawberry mild yellow edge virus Taxonomic position Genus: Potexvirus

(SMYEV)

Family: Alphaflexiviridae

Fragaria spp. (Strawberry)

1024

Geographical distribution SMYEV infection in plants of Fragaria spp. was reported from North America, South America, Europe, Israel, South Africa, Mexico, Australia, New Zealand, Korea, Japan, Canada, and Chile (Spiegel et al. 1986; Converse et al. 1987; Jelkmann et al. 1990; Hepp and Martin 1992; Spiegel and Martin 1992; Cardoni et al. 2002; Martin and Tzanetakis 2006; Cho et al. 2011; Li and Yang 2011a; Rojas et al. 2013; Silva-Rosales et al. 2013; Tzanetakis and Martin 2013, 2014; Ma et al. 2015). Symptoms and host(s) Most strawberry cultivars are symptomless carriers of SMYEV. The wild species F. virginiana, F. vesca, and F. chiloensis show symptoms, and F. ovalis is a symptomless carrier. The following Fragaria species exhibited the following symptoms when inoculated through strawberry aphid vectors: F. vesca – leaflets cupped, chlorotic margins, and vigor reduced. An isolate from the Pacific Coast of North America causes chlorotic vein netting and necrosis of youngest leaves. F. virginiana – leaflets cupped and vigor reduced. Transmission The virus is transmitted by strawberry aphids, Chaetosiphon fragaefolii, C. thomasi, C. jacobi, in a persistent manner, possibly due to the presence of a helper virus (Martin and Tzanetakis 2006). The virus is not transmissible by mechanical inoculation. The virus is not transmissible by contact between plants nor by seed and pollen. The virus-infected planting material is the most efficient dissemination route for virus spread. The virus is graft-transmissible. Virion properties and genome The virions are flexuous filaments, with a modal length of 482 and 13 nm diameter. The virions contain a single, linear molecule of positive-sense ssRNA of 5966 nt (D12517 = NC_003794) and comprise five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type and 18–27 kDa in size (Martin and Converse 1985; Jelkmann et al. 1992; Thompson and Jelkmann 2004; Adams et al. 2004; Cho et al. 2011).

Strawberry mottle virus Taxonomic position Genus: Unassigned

(SMoV)

Family: Secoviridae

Geographical distribution SMoV infection in plants of Fragaria spp. was probably distributed worldwide where strawberries are grown (Mellor and Krczal 1987; Cardoni et al. 2002; Thompson and Jelkmann 2003; Martin and Tzanetakis 2006; Ward et al. 2008; Rojas et al. 2013; Tzanetakis and Martin 2013, 2014). Symptoms and host(s) The virus is symptomless in most of the commercial strawberry cultivars, but in some cultivars the virus-infected strawberry plants have reduced vigor and fruit yield. Because of innumerable strains or variants of SMoV a range of symptoms are produced on F. vesca from barely discernible mottle to severe degeneration.

Fragaria spp. (Strawberry)

1025

Transmission The virus is transmitted by the aphid vectors, Chaetosiphon fragaefolii, C. thomasi, C. minor, C. jacobi, Myzus ascalonicus, M. ornatus, and Aphis gossypii, in a semi-persistent manner (Frazier 1968). The virus is lost by the vector when it molts, does not multiply in the vector, and is not transmitted congenitally to the progeny of the vector. The virus is mechanically sap-transmissible to a number of hosts and also transmissible by grafting. SMoV is not transmissible by contact between plants and not transmitted by seed. Virion properties and genome The virions are isometric and non-enveloped and are 37 nm in diameter (Yoshikawa and Converse 1991). The genome is bipartite with a single-stranded positive-sense RNAs: RNA1 consists of 7036 nt (AJ311875 = NC_003445) and RNA 2 of 5619 nt (AJ311876 = NC_003446) excluding the poly-A tail (Thompson et al. 2002; Thompson et al. 2017).

Strawberry necrotic shock virus Taxonomic position Genus: Ilarvirus

(SNSV)

Family: Bromoviridae

Geographical distribution SNSV infection in plants of Fragaria spp. was first identified in the 1950s by Jorgensen (1957) and later by Frazier et al. (1962) and Converse (1969), and for many years, this disease was thought to be caused by a strain of TSV (Stenger et al. 1987). Strawberry necrotic shock disease is now known to be caused by SNSV, and not by a strain of TSV (Tzanetakis et al. 2004c). SNSV infection in plants of Fragaria spp. was reported from the USA, Australia, Mexico, North America, Europe, Asia, China, New Zealand, and Israel (Tzanetakis et al. 2001; Tzanetakis 2004; Martin and Tzanetakis 2006, 2013; Moyer et al. 2010; Li and Yang 2011b; Sharman et al. 2011; SilvaRosales et al. 2013; Tzanetakis and Martin 2013, 2014). Symptoms and host(s) Irregular distribution of SNSV was found in roots and shoots of strawberry plantlets grown in vitro (Spiegel et al. 1995). The virus-infected plants do not exhibit any external symptoms. The virus has a wide host range, and hosts such as bean, clover, tomato, and weed species, among others, can serve as sources of inoculum. Transmission The virus is spread through seed and pollen and by thrips (Johnson et al. 1984; Kaiser et al. 1982). There is a virus-vector-pollen host relationship, and the transmission of SNSV occurs when feeding thrips cause wounds on plant tissues. Thrips mandibles pierce infective pollen grains and so transfer virus from the pollen grain to the damaged cell (Sdoodee and Teakle 1993). The virus is mechanically sap-transmissible. Virion properties and genome The virions are quasi-isometric particles about 30 nm in diameter. The genome is a tripartite RNA: RNA1 consists of 3429 nt (DQ318818 = NC_008708), RNA2 of 2876 nt (AY743591 = NC_008707), and RNA3 of 2245 nt (AY363228 = NC_008706) (Tzanetakis et al. 2010; Scott 2011a, b).

F

Fragaria spp. (Strawberry)

1026

Strawberry pallidosis-associated virus Taxonomic position Genus: Crinivirus

(SPaV)

Family: Closteroviridae

Geographical distribution For the first time in the 1950s, SPaV was identified in Fragaria spp. in the USA (Frazier and Stubbs 1969) and is present in North America, South America, New Zealand, Australia, Canada, Egypt, China, and the USA (Fulton 1987a; Spiegel and Martin 1998; Tzanetakis 2004, 2010; Tzanetakis et al. 2004a, b, 2013; Martin and Tzanetakis 2006, Martin and Tzanetakis 2013; Wintermantel et al. 2006; Ragab et al. 2009; Constable et al. 2010; Silva-Rosales et al. 2013; Tzanetakis and Martin 2014; Ding et al. 2017; Shi et al. 2018). Symptoms and host(s) The virus-infected strawberry plants will show either latent infection or very mild, non-diagnostic symptoms. Runners are pale and shortened. When infected by severe isolates, some plants may die. Symptoms are most severe in greenhouse-grown plants during the winter. During the summer, symptoms fade unless plants are heavily shaded (Converse and Volk 1990). Transmission The virus is transmitted by the glasshouse whitefly vector, Trialeurodes vaporariorum, in a semipersistent manner (Tzanetakis et al. 2006b). The virus is neither mechanically sap-transmissible nor seed-borne. The virus is graft-transmissible, and the use of runners from infected plants for planting is responsible for most virus spread. Virion properties and genome The virions are non-enveloped, bipartite filamentous particles about 650–850 nm and 700–900 nm in length, and 10–13 nm in diameter. The genome consists of two molecules of positive-sense ssRNA. RNA1 of 8066 nt (NC_005895) and RNA2 of 7978 nt (NC_005896) (Yoshikawa and Converse 1990; Tzanetakis et al. 2005; Kreuze 2011).

Strawberry pseudo mild yellow edge virus Taxonomic position Genus: Carlavirus

(SPMYEV)

Family: Betaflexiviridae

Geographical distribution SPMYEV infection in plants of Fragaria spp. was reported from the USA and Asia (Frazier 1966, 1987; Martin and Tzanetakis 2006; Tzanetakis and Martin 2013). Symptoms and host(s) The virus-infected strawberry cultivars are symptomless. On F. vesca, older leaves become mottled, may exhibit vein yellowing or stippling, later turn necrotic, and die prematurely.

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Transmission The virus is transmitted by the aphid vectors, Chaetosiphon fragaefolii and Aphis gossypii, in a nonpersistent manner. The virus does not multiply in the vector and is lost when the vector molts. The virus is not transmitted congenitally to the progeny of the vector. The virus is mechanically sap-transmissible and also graft-transmissible. Virion properties and genome The virions are rod-shaped particles, 625  12 nm (Yoshikawa and Inouye 1986; Yoshikawa et al. 1986a). The genome consists of a single molecule of linear ssRNA 7.4–7.9 kb in size and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Adams et al. 2004).

Strawberry vein banding virus Taxonomic position Genus: Caulimovirus

(SVBV)

Family: Caulimoviridae

Geographical distribution SVBV infection in plants of Fragaria spp. was described by Frazier (1955a). The virus occurs in temperate regions of eastern and western North America, South America, Africa, Asia, New Zealand, Canada, Chile, the Czechoslovakia, Germany, Hungary, Ireland, Italy, the USA, former USSR, Australia, Turkey, Europe and Brazil (Schoniger 1958; Morris et al. 1980; Frazier and Morris 1987; Honetslegrova et al. 1995; CABI/EPPO 2002; Cardoni et al. 2002; Martin and Tzanetakis 2006; Ratti et al. 2009; Rojas et al. 2013; Tzanetakis and Martin 2013, 2014; Song et al. 2014; Dickison et al. 2017). Symptoms and host(s) Symptoms of virus-infected strawberry plants initially appear on the youngest developing leaf; there is epinasty of midribs and petioles, a tendency for opposite halves of leaflets to be appressed, and irregular, wavy leaflet margins and slight crinkling of the laminae. Usually, these symptoms are mild and are not all present simultaneously. It is not until the affected leaf expands that vein-clearing, followed by yellow vein-banding of some, or all, of the main and secondary veins, becomes visible. Often, this coloration occurs in scattered discontinuous streaks of varying lengths along the main and secondary veins. Spotting of older leaves and twisting of leaflets may occur (OEPP/EPPO 1978). The virus is known to occur only on Fragaria spp. The main host is Fragaria vesca (wild strawberry). Commercial strawberries may also be infected, but diagnostic symptoms are usually only apparent when strawberry latent C “rhabdovirus” is present simultaneously (EPPO/CABI 1996) Transmission The virus is transmitted by the aphid vectors, Chaetosiphon jacobi, C. fragaefolii, and C. thomasi, in a semi-persistent manner. The aphids can acquire the virus during a 30-min-acquisition access feeding period. Virus does not multiply in the vector nor is it transmitted congenitally to the progeny of the vector. The virus is not transmissible by mechanical sap-inoculation or by contact between plants. No seed and pollen transmission is reported. The virus has been transmissible by grafting and dodder (Frazier 1960).

F

Fragaria spp. (Strawberry)

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Virion properties and genome The virions are isometric, non-enveloped, and 45–50 nm in diameter with no obvious surface structure. The genome is a monopartite, circular, double-stranded DNA of 7876 bp (NC_001725) (Kaname 1975; Stenger et al. 1988; Petrzik et al. 1998; Hohn 2011).

Tobacco mosaic virus

(TMV)

Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

TMV infection in plants of Fragaria spp. was reported from Japan and France (Cornuet and Morand 1960; Kaname and Kishi 1973). There is no known vector for this virus. The virus is transmissible by mechanical sap-inoculation, by grafting and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco necrosis virus D Taxonomic position Genus: Betanecrovirus

(TNV-D)

Family: Tombusviridae

TNV-D infection in plants of Fragaria spp. was first reported from Arkansas (USA) by Fulton (1952). The virus was reported from Italy, Bulgaria, California (USA), the Czech Republic, Lithuania, and Japan (Frazier 1955b; Kaname and Kishi 1973; Komuro et al. 1973; Franova-Honetslegrova et al. 1998a, b; Staniulis et al. 2004; Martin and Tzanetakis 2006). The virus-infected F. vesca plants exhibit symptoms of dwarfing, leaf malformation, necrosis, and mottling (Faccioli 1969). The virus is transmitted by a fungal vector Olpidium brassicae. TNV-D is also mechanically saptransmissible to a number of herbaceous plants such as Nicotiana acaulis, N. benthamiana, and N. occidentalis 37 B (systemic hosts) and Ammobium alatum, N. bigelovii, and Petunia hybrida (local hosts). For more details of TNV-D, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Fragaria spp. occurs worldwide including North America, South America, Europe, Africa, Asia, Australia, Israel, and New Zealand (Stace-Smith and Frazier 1971; Spiegel and Cohen 1985; Stace-Smith et al. 1987; Stenger et al. 1987; Guido Herrera and Arturo Lavin 1998; Hokanson et al. 2000; Tzanetakis et al. 2003b; Tzanetakis and Martin 2013). The virus-infected strawberry plants are usually asymptomatic: some commercial cultivars produce mild transient symptoms. The virus is transmitted by the thrips vectors (Frankliniella occidentalis and Thrips tabaci), the virus is present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is mechanically sap-transmissible. The virus is seed-transmitted up to 35% (Johnson et al. 1984). Pollen transmission is also reported. The virus is graft-transmissible. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Fragaria spp. (Strawberry)

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Tomato black ring virus Taxonomic position Genus: Nepovirus

(TBRV)

Family: Secoviridae

TBRV infection in plants of Fragaria spp. was reported from the USA, Asia, and Europe (Guido Herrera and Arturo Lavin 1998; Cardoni et al. 2002; Martin and Tzanetakis 2006). The virus-infected strawberry cultivars exhibit yellow blotching, ringspots, and crinkling of the leaves and finally leads to stunting and eventual death of plants. The virus is transmitted by the nematode vectors, Longidorus attenuatus and L. elongatus, in a non-persistent manner, and is also transmissible by mechanical sapinoculation (Harrison et al. 1961; Brown and Trudgill 1998). The virus is seed-transmitted in Fragaria x ananassa to the extent of 40% (Lister 1960). For more details of TBRV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Fragaria spp. was first reported from the USA (Frazier et al. 1961). The virus spreads in North America, South America, Europe, Africa, Asia, and New Zealand (Converse 1981; Guido Herrera and Arturo Lavin 1998; Cardoni et al. 2002; Martin and Tzanetakis 2006). The virus-infected F. chiloensis plants are symptomless when infected. Symptoms in cultivated strawberries range from none to dwarfing, reduction in runner production, and occasionally mottled leaves in the spring. In F. vesca the symptoms are generally a mild mosaic and sometimes reddish discoloration of the young petioles and leaves. The virus is transmitted by a nematode vector Xiphinema americanum in a non-persistent manner, and by mechanical sap-inoculation to several herbaceous plants. The virus is seed-transmitted in F. vesca and in cultivated strawberry, often without symptoms (Mellor and Stace-Smith 1963). The virus is also transmissible by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

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Latv J Agron 7:49–54 Stenger DC, Mullin RH, Morris TJ (1987) Characterization and detection of the strawberry necrotic shock isolate of Tobacco streak virus. Phytopathology 77:1330–1337 Stenger DC, Mullin RH, Morris TJ (1988) Isolation, molecular cloning, and detection of Strawberry vein banding virus DNA. Phytopathology 78:154–159 Thompson JR, Jelkmann W (2003) The detection and variation of Strawberry mottle virus. Plant Dis 87:385–390 Thompson JR, Jelkmann W (2004) Strain diversity and conserved genome elements in Strawberry mild yellow edge virus. Arch Virol 149:1897–1909 Thompson JR, Leone G, Lindner JL, Jelkmann W, Schoen CD (2002) Characterization and complete nucleotide sequence of Strawberry mottle virus: a tentative member of a new family of bipartite plant picorna-like viruses. J Gen Virol 83:229–239 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Secoviridae. J Gen Virol 98:529–531 Trudgill DL, Brown DJF, McNamara DG (1983) Methods and criteria for assessing the transmission of plant viruses by longidorid nematodes. Rev Nematol 6:133–141 Tzanetakis IE (2004) Molecular characterization of criniviruses and ilarviruses infecting strawberry. PhD dissertation submitted to Oregon State University, USA. Tzanetakis IE (2010) Emerging strawberry virus and virus-like diseases in the world. In: 21st International conference on virus and other graft transmissible diseases of fruit crops. Juliius Kuhn Archiv 427:41–43. Tzanetakis IE, Martin RR (2004) Complete nucleotide sequence of a strawberry isolate of Beet pseudo-yellows virus. Virus Genes 28:239–246 Tzanetakis IE, Martin RR (2005a) First report of strawberry as a natural host of Apple mosaic virus. Plant Dis 89(4):431 Tzanetakis IE, Martin RR (2005b) Fragaria chiloensis cryptic virus: a new strawberry virus found in Fragaria chiloensis plants from Chile. Plant Dis 89:1241 Tzanetakis IE, Martin RR (2005c) New features in the genus Ilarvirus revealed by the nucleotide sequence of Fragaria chiloensis latent virus. Virus Res 112(1–2):32–37 Tzanetakis IE, Martin RR (2007) Strawberry chlorotic fleck: identification and characterization of a novel Closterovirus associated with the disease. Virus Res 124:88–94 Tzanetakis IE, Martin RR (2008) Strawberry chlorotic fleck disease may be caused by a novel closterovirus. Proc XIth IS on small fruit virus diseases. Acta Hortic 780:21–28 Tzanetakis IE, Martin RR (2013) Expanding field of strawberry viruses which are important in North America. Int J Fruit Sci 13:184–195 Tzanetakis IE, Martin RR (2014) Incidence of major strawberry viruses in North America. Acta Hortic 1049:595–598 Tzanetakis IE, Mackey IC, Martin RR (2001) Strawberry necrotic shock virus: a new virus previously throught to be Tobacco streak virus. Acta Hortic 656:27–31 Tzanetakis IE, Wintermantel WM, Martin RR (2003a) First report of Beet pseudo yellows virus in strawberry in the United States: a second crinivirus able to cause pallidosis disease. Plant Dis 87:1398 Tzanetakis IE, Mackey IC, Martin RR (2003b) Tobacco streak virus in strawberry. An old problem, a new virus? Phytopathology 93:S85 Tzanetakis IE, Halgren AB, Wintermantel WM, Keller KE, Martin RR (2004a) Two Criniviruses are associated with the strawberry pallidosis disease. Acta Hortic 656:21–26 Tzanetakis IE, Halgren AB, Keller KE, Hokanson SC, Mass JL, McCarthy PL, Martin RR (2004b) Identifiatin and detection of a virus associated with strawberry pallidosis disease. Plant Dis 88:383–390 Tzanetakis IE, Mackey IC, Martin RR (2004c) Strawberry necrotic shock virus is a distinct virus and not a strain of Tobacco streak virus. Arch Virol 149:2001–2011 Tzanetakis IE, Reed J, Martin RR (2005) Nucleotide sequence, genome organization and phylogenetic analysis of Strawberry pallidosis-associated virus, a new member of the genus Crinivirus. Arch Virol 150:273–286 Tzanetakis IE, Postman JD, Gergerich RC, Martin RR (2006a) A virus between families: nucleotide sequence and evolution of Strawberry latent ringspot virus. Virus Res 121:199–204 Tzanetakis IE, Wintermantel WM, Cortez AA, Barnes JE, Barrett SM, Bolda MP, Martin RR (2006b) Epidemiology of strawberry pallidosis-associated virus and occurrence of pallidosis disease in North America. Plant Dis 90(10):1343–1346

F

Franklinia alatamaha (Franklin tree)

1034

Tzanetakis IE, Price R, Martin RR (2008) Nucleotide sequence of the tripartite Fragaria chiloensis cryptic virus and presence of the virus in the Americas. Virus Genes 36:267–272 Tzanetakis IE, Martin RR, Scott SW (2010) Genomic sequences of Blackberry chlorotic ringspot virus and Strawberry necrotic shock virus and the phylogeny of viruses in subgroup 1 of the genus Ilarvirus. Arch Virol 155:557–561 Tzanetakis IE, Martin RR, Wintermantel WM (2013) Epidemiology of Criniviruses: an emerging problem in world agriculture. Front Microbiol 4:119. https://doi.org/10.3389/fmicb.2013.00119 Vainio EJ, Chiba S, Ghabrial SA, Maiss E, Roossinck M, Sabanadzovic S, Suzuki N, Xie J, Nibert M, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Partitiviridae.J Gen Virol 99:17–18 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Rhabdoviridae. J Gen Virol 99:447–448 Ward LI, Quinn BD, Tang J, Wei T, Clover GRG (2008) Strawberry mottle virus detected on strawberry in New Zealand. Plant Pathol 57:1172 Wetzel T, Meunier L, Jaeger U, Reustle GM, Krczal G (2001) Complete nucleotide sequences of the RNAs 2 of German isolates of grapevine fanleaf and Arabis mosaic nepoviruses. Virus Res 75(2):139–145 Wetzel T, Beck A, Wegener U, Krczal G (2004) Complete nucleotide sequence of the RNA 1 of a grapevine isolate of Arabis mosaic virus. Arch Virol 149(5):989–995 Wintermantel WM, Fuentes S, Chuquillanqui C, Salazar LF (2006) First report of Beet pseudo-yellows virus and Strawberry pallidosis associated virus in strawberry in Peru. Plant Dis 90:1457 Yoshikawa N, Converse RH (1990) Strawberry pallidosis disease: distinctive dsRNA species associated with latent infections in indicators and in diseased strawberry cultivars. Phytopathology 80:543–548 Yoshikawa N, Converse RH (1991) Purification and some properties of Strawberry mottle virus. Ann Appl Biol 118:565–576 Yoshikawa N, Inouye T (1986) Purification, characterization and serology of Strawberry pseudo mild yellow-edge virus. Ann Phytopathol Soc Japan 52:643–652 Yoshikawa N, Inouye T (1988) Strawberry viruses occurring in Japan. Acta Hortic 236:59–67 Yoshikawa N, Poolpol P, Inouye T (1986a) Use of dot immunobinding assay for rapid detection of strawberry pseudo mild yellow-edge virus. Ann Phytopathol Soc Japan 52:728–731 Yoshikawa N, Inouye T, Converse RH (1986b) Two types of rhabdovirus in strawberry. Ann Phytopathol Soc Japan 52:437–444 Zeller SM, Vaughan EK (1932) Crinkle disease of strawberry. Phytopathology 22:709–713

Franklinia alatamaha (Franklin tree) Family: Theaceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Franklinia alatamaha was reported from Georgia, USA (Ruter and Gitaitis 1993). The virus-infected franklin trees do not exhibit any symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

References Ruter JM, Gitaitis RD (1993) Impatiens necrotic spot virus in woody landscape plants in Georgia. Plant Dis 77:318

Fraxinus spp. (Fraxinus americana; F. excelsior) (White ash; Common ash)

1035

Fraxinus spp. (Fraxinus americana; F. excelsior) (White ash; Common ash) Family: Oleaceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Trees/Shrubs

(ArMV)

Family: Secoviridae

ArMV infection in plants of Fraxinus excelsior was reported from the UK (Cooper 1975; Cooper and Sweet 1976). The virus-infected common ash trees exhibit symptoms of chlorotic mottling, chevrons, or oak leaf patterns. The virus is transmitted by the nematode vector, Xiphinema diversicaudatum, in a non-persistent manner, and is also transmissible by mechanical sap-inoculation, and by grafting. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Cherry leaf roll virus Taxonomic position Genus: Nepovirus

(CLRV)

Family: Secoviridae

CLRV infection in plants of Fraxinus excelsior was reported from Germany and Finland (Hamacher and Quadt 1991; Rebenstorf et al. 2006; von Bargen et al. 2009). The virus-infected common ash plants exhibit deformed leaves, chlorotic ringspots, and line pattern symptoms on leaves. The virus is transmitted by nematode vectors in a non-persistent, manner and is also transmissible by mechanical sap-inoculation, and grafting. For more details of CLRV, refer to Prunus avium.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV infection in plants of Fraxinus americana was reported from New York, USA (Lana and Agrios 1973, 1974; Castello et al. 1984). The virus-infected white ash plants exhibit symptoms of line patterns, mosaic, chlorotic spots, and mottling. There is no known vector for this virus. The virus is transmissible by mechanical sap-inoculation, transmitted by grafting, and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

F

Fraxinus spp. (Fraxinus americana; F. excelsior) (White ash; Common ash)

1036

TRSV infection in plants of Fraxinus americana was reported from the northern states of USA (Hibben and Bozarth 1972; Castello et al. 1984). The virus-infected white ash plants exhibit symptoms of chlorotic spots, rings, vein-banding patterns, red colored spots, and rings. The virus is transmitted by the nematode vector, Xiphinema rivesi, in a non-persistent manner, and is also transmissible by mechanical sap-inoculation (Hibben and Walker 1971). The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato bushy stunt virus Taxonomic position Genus: Tombusvirus

(TBSV)

Family: Tombusviridae

TBSV infection in plants of Fraxinus excelsior was reported by Novak and Lanzova (1980). The virus is transmissible by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants. There is no known vector for this virus. For more details of TBSV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Fraxinus americana was reported from New York, USA (Ferris and Castello 1988; Hibben et al. 1988). The virus-infected common ash plants exhibit symptoms of chlorotic spots, blotches, and line patterns. The virus is transmitted by nematode vectors in a nonpersistent manner, and is also transmissible by mechanical sap-inoculation, and by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

White ash mosaic virus

(WAMV)

Taxonomic position WAMV is a tentative member of the family Betaflexiviridae. Geographical distribution WAMV infection in plants of Fraxinus spp. was reported from Minnesota, Ohio, New York, and Illinois, USA (Machado-Caballero et al. 2013). Symptoms and host(s) The virus-infected white ash plants exhibit mosaic and decline symptoms. The virus naturally infects green F. pennsylvanica. Transmission The virus is mechanically sap-transmissible but not through seed or by the aphid, Myzus persicae.

Freesia spp.

1037

Virion properties and genome The virions are flexuous filaments 720 nm in length. The genome is a single positive-sense, singlestranded RNA of 9.0 kb (GU906791 and DQ412998) (Machado-Caballero et al. 2013).

References Castello JD, Amico LA, O'Shea MT (1984) Detection of Tobacco mosaic and Tobacco ringspot viruses in white ash trees by enzyme-linked immunosorbent assay. Plant Dis 68:787–790 Cooper JI (1975) Arabis mosaic virus, a cause of chlorotic symptoms in leaves of Fraxinus excelsior L. Plant Pathol 24:114–116 Cooper JI, Sweet JB (1976) The detection of viruses with nematode vectors in six woody hosts. Forestry 49:73–78 Ferris MA, Castello JD (1988) Detection of Tomato ringspot virus in white ash and adjacent vegetation in central New York. Can J Forest Res 18:813–817 Hamacher J, Quadt A (1991) Light- and electron microscopic studies of Cherry leaf roll virus (CLRV) on European Ash (Fraxinus excelsior L.). J Phytopathol 131:215–226 Hibben CR, Bozarth RF (1972) Identification of an ash strain of Tobacco ringspot virus. Phytopathology 62:1023–1029 Hibben CR, Walker JT (1971) Nematode transmission of the ash strain of Tobacco ringspot virus. Plant Dis Reptr 55:475–478 Hibben CR, Reese J, Castello JD (1988) Identification of Tomato ringspot virus in ash in New York. Plant Dis 72:175 Lana AO, Agrios GN (1973) A disease of white ash caused by Tobacco mosaic virus. Phytopathology 63:203 Lana A, Agrios G (1974) Properties of a Strain of Tobacco mosaic virus isolated from white ash trees. Phytopathology 64:1490–1495 Machado-Caballero JE, Lockhart BE, Mason SL, Mollov D, Smith JA (2013) Identification, transmission, and partial characterization of a previously undescribed flexivirus causing a mosaic disease of ash (Fraxinus spp.) in the USA. Online. Plant Health Prog. https://doi.org/10.1094/PHP-2013-0509-01-RS Novak JB, Lanzova J (1980) Demonstration of Tomato bushy stunt virus in some forest tree species and plants. Lesnictvi (Prague) 26:1009–1016 Rebenstorf K, Candresse T, Dulucq JM, Buttner C, Obermeier C (2006) Host species-dependent population structure of a pollen-borne plant virus, Cherry leaf roll virus. J Virol 80:2453–2462 von Bargen S, Grubits E, Jalkanen R, Büttner C (2009) Cherry leaf roll virus – an emerging virus in Finland? Silva Fenn 43:727–738

Freesia spp. Family: Iridaceae

Ornamental

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV occurs in plants of Freesia spp. wherever the crop is grown (Bellardi and Bertaccini 1989; Nemethy 1994; Kumar et al. 2009b). The virus-infected freesia plants usually have leaves which are chlorotic, and corm necrosis in some cultivars is occasionally observed. The flowers show flower break and often have distorted pistils and stamens. The virus is transmitted by the aphid species such as Myzus persicae and Macrosiphum euphorbiae in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

F

Freesia spp.

1038

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV has been reported in plants of Freesia spp. in Italy, Hungary, and several other countries where the crop is grown (Bellardi and Bertaccini 1989; Mondelli and Rana 1985; Nemethy 1994). The virusinfected freesia plants are stunted and produce chlorotic leaves. The flowers exhibit flower break and distortion. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Freesia leaf necrosis virus Taxonomic position The virus is a tentative member of the genus Varicosavirus and family Rhabdoviridae. Geographical distribution The virus was first reported in plants of Freesia refracta from the Netherlands by Van Dorst (1973). The virus spreads in Germany, Ireland, Italy, the Netherlands, and the United Kingdom (Bouwen 1994; Meekes and Verbeek 2011). Symptoms and host(s) The virus-infected freesia plants exhibit symptoms of chlorotic flecks and spots in leaves, which become necrotic. Flowers and corms are unaffected. Transmission The virus is transmitted by the fungal vector, Olpidium brassicae (Van Dorst and Peters 1988). The virus is transmissible by mechanical sap-inoculation to less than three plant families. The virus is not transmitted by seed. Virion properties and genome The virions are labile structures of c. 21–25 nm wide and 160–390 nm long in the compact form; loose spiral structures of similar width but much greater length are also observed.

Freesia mosaic virus Taxonomic position Genus: Potyvirus

(FreMV)

Family: Potyviridae

Geographical distribution FreMV infection in plants of Freesia refracta was first reported from the Netherlands (Van Koot et al. 1954). The virus occurs in Australia, Ireland, Korea, the Netherlands, New Zealand, Hungary, Italy, the United Kingdom, and the USA (Bellardi and Bertaccini 1989; Nemethy 1994; Kumar et al. 2009a; Pham et al. 2011).

Freesia spp.

1039

Symptoms and host(s) The virus-infected freesia plants are generally symptomless, and in some cultivars, flowers are “broken” or mottled but not distorted. Sometimes conspicuously chlorotic leaves are also observed (Van Koot et al. 1954). Transmission Aphid vectors, such as Macrosiphum euphorbiae and Myzus persicae, transmit the virus in a nonpersistent manner, and is also transmissible by mechanical sap-inoculation (Kumar et al. 2009a). Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9489 nt (FM206346 = NC_014064; GU214748) (Choi et al. 2010; Revers and Garcia 2015; Wylie et al. 2017).

Freesia sneak ophiovirus Taxonomic position Genus: Ophiovirus

(FreSV)

Family: Aspiviridae

Geographical distribution FreSV infection in plants of Freesia spp. has been reported from the Netherlands, Italy, Bulgaria, Korea, and Virginia (Vaira et al. 2006, 2009, 2015; Hansen et al. 2009; Meekes and Verbeek 2011; Bobev et al. 2013; Jeong et al. 2014). Symptoms and host(s) The virus-infected freesia plants exhibit initial symptoms consisting of scattered pale, chlorotic, and interveinal lesions that coalesce. Later, irregular brown to black necrotic blotches partially cover the leaves. Flower break is also observed. Transmission The virus is transmitted by the soil-borne fungus Olpidium brassicae. The virus is also mechanically sap transmissible. Virion properties and genome The virions are non-enveloped, naked filamentous nucleocapsids about 3 nm in diameter, forming kinked circles of at least two different contour lengths (300–500 and 1500–2500 nm). The circles (open form) can collapse to form pseudo-linear duplex structures of 9–10 nm in diameter (collapsed form). The genome is a linear, negative-sense, single-stranded RNA and consists of three segments. RNA3 contains 1456 nt (DQ885455). Additional partial coat protein sequences are available (e.g. FJ807730) (Vaira et al. 2009; Milne et al. 2011; Garcia 2012).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(INSV)

F

Freesia spp.

1040

INSV infection in plants of Freesia spp. was reported from New Zealand (Lebas et al. 2004; Elliott et al. 2009). The virus-infected freesia plants exhibit symptoms of chlorotic and necrotic leaf streaking. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in plants of Freesia spp. was reported from Hungary and the Netherlands (Asjes 1989, 1994; Nemethy 1994). The virus-infected freesia plants are stunted, flower stems have fewer and smaller flowers, roots are poorly developed, and corm yield is much reduced. The virus is transmitted by the nematode vectors, viz., Trichodorus and Paratrichodorous, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

References Asjes CJ (1989) Tobacco rattle virus and other soil-borne viruses in ornamental bulb culture in The Netherlands. Ann Appl Biol 22:83–86 Asjes CJ (1994) Occurrence of Tobacco rattle virus in ornamental bulbous crops in the Netherlands. Acta Hortic 377:349–356 Bellardi MG, Bertaccini A (1989) Virus diseases of Freesia in Italy. Adv Hortic Sci 3:29–32 Bobev SG, Taphradjiiski OI, Hammond J, Vaira AM (2013) First report of Freesia sneak virus associated with foliar necrosis of Freesia refracta in Bulgaria. Plant Dis 97:1514 Bouwen I (1994) Freesia leaf necrosis: some of its mysteries revealed. Acta Hortic 377:311–318 Choi HI, Lim HR, Song YS, Kim MJ, Choi SH, Song YS, Bae SC, Ryu KH (2010) The complete genome sequence of Freesia mosaic virus and its relationship to other potyviruses. Arch Virol 155:1183–1185 Elliott DR, Lebas BSM, Ochoa-Corona FM, Tang J, Alexander BJR (2009) Investigation of Impatiens necrotic spot virus outbreaks in New Zealand. Aust Plant Pathol 38:490–495 Garcia ML (2012) Ophioviruses: State of the art. In: Garcia M, Romanowski V (eds) Viral genomes-Molecular structure, diversity, gene expression mechanisms and host-virus interactions. Intech Open Limited, London ISBN:978-953-510098-0. https://doi.org/10.5772/29707 Hansen M, Vaira AM, Murphy C, Hammond J, Dienelt M, Bush E, Sutula C (2009) Freesia sneak virus (FreSV) on freesia: a first detection for Virginia and the United States. Department of Plant Pathology, Physiology and Weed Science, VPI&SU, Blacksburg, p 1 Jeong M, Choi YJ, Joa JH, Choi KS, Chung BN (2014) First report of Freesia sneak virus in commercial Freesia hybrida cultivars in Korea. Plant Dis 98:162 Kumar Y, Hallan V, Zaidi AA (2009a) First finding of Freesia mosaic virus infecting freesia in India. Plant Pathol 58:404 Kumar Y, Hallan V, Zaidi AA (2009b) Identification and characterization of Bean yellow mosaic virus infecting Freesia. J Plant Biochem Biotechnol 18:253–255 Lebas BSM, Ochoa-Corona FM, Elliott DR, Tang Z, Alexander BJR, Froud KJ (2004) An investigation of an outbreak of Impatiens necrotic spot virus in New Zealand. Phytopathology 94:S57 Meekes ETM, Verbeek M (2011) New insights into Freesia leaf necrosis disease. Acta Hortic 901:231–236 Milne RG, Garcia ML, Vaira AM (2011) Ophiovirus. Ophioviridae. In: The Springer index of viruses. Springer, New York, pp 995–1003. https://doi.org/10.1007/978-0-387-95919-1_155 Mondelli D, Rana GL (1985) II virus del mosaico del cetriolo su freesia in Peglia. Inf Fitopatol 35:41–42 Nemethy IZ (1994) Survey on virus diseases of bulbous flowers in Hungary. Acta Hortic 377:267–274 Pham KTK, de Kock MJD, Lemmers MEC, Derks AFLM (2011) Molecular identification of potyviruses infecting bulbous ornamentals by the analysis of coat protein (CP) sequences. Acta Hortic 901:167–172 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Vaira AM, Lisa V, Costantini A, Masenga V, Rapetti S, Milne RG (2006) Ophioviruses infecting ornamentals and a probable new species associated with a severe disease in freesia. Acta Hortic 722:191–200

Fritillaria thumbergii (Thunberg fritillary)

1041

Vaira AM, Hansen MA, Murphy C, Reinsel MD, Hammond J (2009) First report of Freesia sneak virus in Freesia sp. in Virginia. Plant Dis 93:965 Vaira A, Vallino M, Lenzi R, Masenga V, Lisa V, Constatini A, Salvi D, Carra A, Hammond J (2015) Detection of mixed virus population in freesia plants with necrotic disease. Acta Hortic 1072:173–178 Van Dorst HJM (1973) Two new disorders in freesias. Neth J Plant Pathol 79:130–137 Van Dorst HJM, Peters D (1988) Experiences with the freesia leaf necrosis agent and its presumed vector, Olpidium brassicae. In: Cooper JI, Asher MJC (eds) Developments in applied biology 2, viruses with fungal vectors. AAB Wellesbourne, UK, pp 315–322 Van Koot Y, van Slogteren DHM, Cremer MC, Camfferman J (1954) Tijdschr Plziekt 60:157 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354

Fritillaria thumbergii (Thunberg fritillary) Family: Liliaceae

Medicinal

Fritillary virus Y

(FVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution FVY infection in plants of Fritillaria thumbergii was reported from China (Chen et al. 2006). Symptoms and host(s) The virus-infected thunberg fritillary plants exhibit mottling symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also transmissible mechanically to the original host but not to any of a range of commonly used indicator plants. No local lesion host was identified that would enable the virus to be propagated independently. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome is a single molecule of positive-sense, single-stranded RNA of 9656 nt (AM039800 = NC_010954) (Chen et al. 2006; Revers and Garcia 2015; Wylie et al. 2017).

Potato leafroll virus Taxonomic position Genus: Polerovirus

(PLRV)

Family: Luteoviridae

PLRV infection in plants of Fritillaria thumbergii was reported from China (Tu et al. 2006). The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical inoculation. For more details of PLRV, refer to Solanum tuberosum.

F

Fuchsia spp.

1042

Thunberg fritillary mosaic virus Taxonomic position Genus: Potyvirus

(TFMV)

Family: Potyviridae

Geographical distribution TFMV-infected in plants of Fritillaria thumbergii were reported from China (Wei et al. 2005). Symptoms and host(s) The virus-infected thunberg fritillary plants exhibit mottle and mosaic symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also transmissible mechanically to the original host but not to any of a range of commonly used indicator plants. No local lesion host was identified that would enable the virus to be propagated independently. Virion properties and genome The virions are non-enveloped, flexuous filaments, 750 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 9723 nt (AJ851866 = NC_007180) (Wei et al. 2005; Revers and Garcia 2015; Wylie et al. 2017).

References Chen J, Zheng H-Y, Shi Y-H, Adams MJ, Wei C-B, Lin L, Chen J-P (2006) Detection and characterisation of a second potyvirus from Thunberg fritillary in China. Arch Virol 151:439–447 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Tu S, Qin C, Chen C (2006) Fritillaria thumbergii – a new host of Potato leafroll virus in China. Aust Plant Dis Notes 1:31–32 Wei CB, Chen J, Zhang QY, Shi YH, Lin L, Zheng HY, Adams MJ, Chen JP (2005) A new potyvirus from Thunberg fritillary (Fritillaria thunbergii Miq.) in Zhejiang, China. Arch Virol 150:1271–1280 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354

Fuchsia spp. Family: Onagraceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Fuchsia spp. was reported from Italy, France, and Minnesota, USA (Roggero et al. 2000; Lockhart 2005). The virus-infected fuchsia plants exhibit severe mosaic, leaf mottling, and leaf deformation symptoms. The virus is transmitted by a number of aphid vectors in a

Fuchsia spp.

1043

non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of host plants. For more details of CMV, refer to Cucumis sativus.

Fuchsia latent virus

(FLV)

Taxonomic position FLV is a tentative member of the genus Carlavirus and family Betaflexiviridae. Geographical distribution FLV was first reported in plants of Fuchsia speciosa  F. hybrida from British Columbia, Canada, by Johns et al. (1980). The virus spreads in Belgium, Italy, and New Zealand (Perez-Egusquiza et al. 2011). Symptoms and host(s) The virus-infected fuchsia plants are typically symptomless. Some cultivars exhibit symptoms of mild chlorosis, mild mottle, and purple leaf spotting. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically saptransmissible to less than three plant families. Virion properties and genome The virions are flexuous filaments, 650 nm in length, and 12 nm wide. Partial sequences of several isolates are available (e.g. HQ197676) (Perez-Egusquiza et al. 2011).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Fuchsia spp. has been recorded by Wick and Dicklow (1999). The virusinfected fuchsia plants exhibit symptoms of foliar ringspots and necrotic patches. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV infection in plants of Fuchsia spp. was reported from the UK (Harju et al. 2011). There is no known vector for this virus. The virus is mechanically sap-transmissible and is also transmissible by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

F

Furcraea spp.

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Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Fuchsia spp. was reported from Canada, Portugal, and Greece (Tehrani et al. 1990; Louro 1996; Chatzivassiliou et al. 2000). The virus-infected fuchsia plants exhibit expanding, necrotic, concentric ringspot symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Chatzivassiliou E, Livieratos I, Jenser G, Katis N (2000) Ornamental plants and thrips populations associated with Tomato spotted wilt virus in Greece. Phytoparasitica 28:257–264 Harju VA, Skelton AL, Monger WA, Forde SMD, Daly M, Nixon T, Lawson R, Bennett S, Mumford RA (2011) A report of viruses, viroids, and phytoplasmas found in ornamental plants from 1999–2007: findings from Central Science Laboratory, UK. Acta Hortic 901:223–229 Johns LJ, Stace-Smith R, Kadota DY (1980) Occurrence of a rod-shaped virus in fuchsia cultivars. Acta Hortic 110:195–204 Lockhart BEL (2005) Three previously unrecorded viral diseases of Astilbe, Fuchsia, and Thermopsis species in Minnesota. Plant Dis 89:775 Louro D (1996) Detection and identification of tomato spotted wilt virus and impatiens necrotic spot virus in Portugal. Acta Hortic 431:99–105 Perez-Egusquiza Z, Liefting LW, Veerakone V, Clover GRG, Ciuffo M (2011) First report of a carlavirus in Fuchsia spp. in New Zealand. Plant Dis 95:1484 Roggero P, Ciuffo M, Lenzi R (2000) Natural infection by Cucumber mosaic virus (Cucumovirus; Bromoviridae) in Fuchsia boliviana Carr and the insusceptibility of other Fuchsia species and hybrids to the virus. Plant Pathol 49(6):802 Tehrani B, Allen WR, Matteoni JA (1990) Update on the incidence of tomato spotted wilt virus in greenhouses. Can. Plant Dis Surv 70:102–103 Wick R, Dicklow B (1999) Diseases in fuchsia. Common names of plant diseases. Online publication. The American Phytopathological Society, St. Paul

Furcraea spp. Family: Asparagaceae

Fiber crop

Furcraea necrotic streak virus Taxonomic position Genus: Macanavirus

(FNSV)

Family: Tombusviridae

Geographical distribution FNSV infection was first reported in plants of Furcraea spp. from Colombia by Galvez et al. (1976). The virus spreads in Colombia and South America (Dabek and Castano 1978; Morales et al. 1992).

Furcraea spp.

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Symptoms and host(s) The virus-infected furcraea plants exhibit symptoms of chlorotic and then necrotic streaks. Transmission The virus may have a soil-inhabiting vector. The virus is not transmitted by either Planococcus citri or Saissetia caffea. The virus is transmissible by mechanical inoculation to less than three families and also by grafting. Virion properties and genome The virions are isometric, 28 nm in diameter, and have T = 3 icosahedral symmetry. The genome is a monopartite, linear, single-stranded, positive-sense RNA of 3966 nt (FJ768020 = NC_020469) (Morales et al. 1992).

References Dabek AJ, Castano JJ (1978) The occurrence, symptomatology, transmission and virus aetiology of macana disease of fique (Furcraea spp.) in Colombia, South America. Phytopathol Z 92:57–69 Galvez GE, Castano M, Vesga B, Kitajima EW (1976) Purification and serology of the necrotic streak mosaic virus of sisal. Proc Am Phytopathol Soc 3:296 Morales F, Castano M, Calvert L, Arroyave JA (1992) Furcraea necrotic streak virus: an apparent new member of the dianthovirus group. J Phytopathol 134:247–254

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G

Gaillardia spp. Family: Asteraceae

Ornamental

Bidens mottle virus Taxonomic position Genus: Potyvirus

(BiMoV)

Family: Potyviridae

BiMoV infection in plants of Gaillardia grandiflora was reported from Florida (USA) (Logan et al. 1984). The virus-infected gaillardia plants do not show apparent symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BiMoV, refer to Bidens spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV was reported from Lithuania in plants of Gaillardia spp. and this virus is worldwide in distribution (Samuitiene and Navalinskiene 2008; Valiunas et al. 2008). The virus-infected gaillardia plants exhibit mosaic symptoms on leaves and flower breaking. Petals are distorted and display variously shaped and colored spots and streaks. Plants are generally stunted. The virus is transmitted by a large number of aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Gaillardia latent virus Taxonomic position Genus: Carlavirus

(GaiLV)

Family: Betaflexiviridae

© Springer Nature India Private Limited 2019 K. S. Sastry et al., Encyclopedia of Plant Viruses and Viroids, https://doi.org/10.1007/978-81-322-3912-3

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Gaillardia spp.

Geographic distribution GaiLV was isolated from plants of Gaillardia aristata in Germany (Menzel et al. 2015). Symptoms and host(s) The virus-infected gaillardia plants displayed no obvious symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically transmissible to three Nicotiana spp. Virion properties and genome The virions are slightly flexuous filaments, 610–700 nm in length and 12–15 nm in diameter. The genome consists of a single molecule of linear, single-stranded RNA of 8659 nt (NC_023892) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Adams et al. 2004).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Gaillardia spp. was reported from the USA (Wick 2003). The virus-infected gaillardia plants exhibit symptoms of mottling/mosaic, foliar distortion and chlorosis, and ringspots. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Moroccan pepper virus Taxonomic position Genus: Tombusvirus

(MPV)

Family: Tombusviridae

MPV infection in plants of Gaillardia grandiflora was reported from Southern Iran (Alemzadeh and Ghorbani 2016). The virus-infected gaillardia plants exhibit mosaic and ringspot symptoms. The virus is transmissible by mechanical sap-inoculation, and also by grafting. There is no known vector for this virus. For more details of MPV, refer to Capsicum annuum.

Pelargonium leaf curl virus Taxonomic position Genus: Tombusvirus

(PLCV)

Family: Tombusviridae

PLCV infection in plants of Gaillardia grandiflora was reported from Southern Iran (Alemzadeh and Ghorbani 2016). The virus-infected gaillardia plants exhibit leaf curling symptoms. The virus is mechanically sap-transmissible. There is no vector is known for this virus. For more details of PLCV, refer to Pelargonium spp.

Gaillardia spp.

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Tomato leaf curl Bangladesh virus Taxonomic position Genus: Begomovirus

(ToLCBV)

Family: Geminiviridae

ToLCBV infection in plants of Gaillardia spp. was reported from Rajasthan (India) (Mahatma and Mahatma 2012). The virus-infected gaillardia plants exhibit vein yellowing and in extreme cases, the entire leaflets showed severe yellowing, crumpling, and distortion. Infected plants were dwarfed with smaller-sized flowers and had reduced or no commercial value. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. The virus is graft-transmissible. For more details of ToLCBV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

Gaillardia spp. (G. aristata; G. grandiflora; G. pulchella) have been reported to be infected by TSWV in Canada and elsewhere (Tehrani et al. 1990; Cho et al. 1987; Parrella et al. 2003). The virus-infected gaillardia plants exhibit symptoms of dieback of the growing tips and dark streaking of the terminal stems. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Alemzadeh E, Ghorbani A (2016) Occurrence of Pelargonium leaf curl virus and Moroccan pepper virus on natural hosts. Australas Plant Dis 11:8 Cho JJ, Mau RFL, Mitchell WC, Gonsalves D, Yudin LS (1987) Host list of plants susceptible to Tomato spotted wilt virus (TSWV), Research Extension Series 078. Hawaii Institute of Tropical Agriculture and Human Resources, Honolulu Logan AE, Zettler FW, Christie SR (1984) Susceptibility of Rudbeckia, Zinnia, Ageratum, and other bedding plants to Bidens mottle virus. Plant Dis 68:260–262 Mahatma L, Mahatma MK (2012) First report of a variant of Tomato leaf curl Bangladesh virus infecting Gaillardia. New Dis Rep 26:4 Menzel W, Hamacher J, Winter S (2015) Characterization of a New Carlavirus from Gaillardia aristata. Acta Hortic 1072:129–133 Parrella G, Gognalons P, Gebre-Selassi K, Vovlas C, Marchoux G (2003) An update of the host range of tomato spotted wilt virus. J. Plant Pathol 85(4, special issue):227–264 Samuitiene M, Navalinskiene M (2008) Occurrence of Cucumber mosaic cucumovirus on ornamental plants in Lithuania. Zemdirbyste-Agric 95:135–143 Tehrani B, Allen WR, Matteoni JA (1990) Update on the incidence of tomato spotted wilt virus in greenhouses. Can Plant Dis Surv 70:102–103 Valiunas D, Samuitiene M, Navalinskiene M, Davis RE (2008) Identification of viral and phytoplasmal agents causing diseases in Gaillardia Foug. plants in Lithuania. Agron Res 6:109–118 Wick RL (2003) Floriculture: fact sheets: pest management: impatiens Necrotic spot virus and Tomato spotted wilt virus. 4 p

G

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Galanthus spp. (Snowdrop)

Galanthus spp. (Snowdrop) Family: Amaryllidaceae

Medicinal plant

Snowdrop virus Y (SVY) Taxonomic position SVY is a tentative member of the genus Potyvirus and family Potyviridae Geographic distribution SVY infection in plants of Galanthus spp. was reported from the UK (Monger Unpublished – EU927399). Transmission The virus is presumed to be transmitted by aphid vectors. Virion properties and genome Virions are flexuous filaments. The genome is a single molecule of positive-sense single-stranded RNA. A partial polyprotein gene sequence of 1702 nt is available (EU927399) (Wylie et al. 2017).

References Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Galeopsis spp. (Hemp-nettle) Family: Lamiaceae

Weed host

Apple mosaic virus

(ApMV)

Taxonomic position Genus: Ilarvirus

Family: Bromoviridae

ApMV infection in plants of Galeopsis spp. was reported from Turkey (Arli Sokmen et al. 2005). No insect vector is known for this virus. The virus is mechanically sap-transmissible. For more details of ApMV, refer to Malus domestica.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

Galinsoga parviflora (Guasca)

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CMV infection in plants of Galeopsis spp. plants was reported from Italy (Vicchi and Bellardi 1988). The virus-infected hemp-nettle plants exhibit mosaic mottling, chlorosis, and leaf deformation symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

References Arli Sokmen M, Kutluk Yilmaz ND, Mennan H, Sevik MA (2005) Natural weed hosts of Apple mosaic virus in hazelnut orchards in Turkey. J Plant Pathol 87:239–242 Vicchi V, Bellardi MG (1988) The role of weeds in the epidemiology of gladiolus viruses and MLO. Acta Hortic 234:371–378

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Galinsoga parviflora (Guasca) Family: Asteraceae

Galinsoga mosaic virus Taxonomic position Genus: Gallantivirus

Ornamental

(GaMV)

Family: Tombusviridae

Geographical distribution GaMV was first reported in plants of Galinsoga parviflora from Australia by Behncken (1970). The virus spreads in Australia (Skotnicki and Gibbs 1981). Symptoms and host(s) The virus-infected guasca plants exhibit mosaic symptoms. Transmission The virus is transmissible by mechanical inoculation to several three to nine plant families. No vector is identified. The virus is not transmissible by contact between plants and not transmitted by seed (Shukla et al. 1979). Virion properties and genome The virions are isometric and 28–34 nm in diameter and have T = 3 icosahedral symmetry. The genome is a monopartite, linear, single-stranded, positive-sense RNA of 3803 nt (Y13463 = NC_001818) (Skotnicki and Gibbs 1981; Ciuffreda et al. 1998).

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Galinsoga parviflora was reported from Argentina and the Czech Republic (Gracia and Feldman 1989; Mertelík et al. 1996). The virus-infected guasca plants show no symptoms or only a mild mosaic with stunted growth. The virus is transmitted by thrips vectors in a persistent-

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Galium aparine (Cleavers)

propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Behncken GM (1970) Some properties of a virus from Galinsoga parviflora. Aust J Biol Sci 23:497–501 Ciuffreda P, Rubino L, Russo M (1998) Molecular cloning and complete nucleotide sequence of galinsoga mosaic virus genomic RNA. Arch Virol 143(1):173–180 Gracia O, Feldman JM (1989) First report of tomato spotted wilt virus on celery and three weed species in Argentina. Plant Dis 73:859 Mertelík J, Götsová B, Mokrá V (1996) Epidemiological aspects of Tomato spotted wilt virus infection in the Czech Republic. Acta Hortic 432:368–375 Shukla DD, Shanks GJ, Teakle DS, Behncken GM (1979) Mechanical transmission of Galinsoga mosaic virus in soil. Aust J Bio Sci 32:267–276 Skotnicki A, Gibbs A (1981) Some properties of the virions of galinsoga mosaic virus. Aust Plant Pathol 10:27–27

Galium aparine (Cleavers) Family: Rubiaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Medicinal

(CMV)

Family: Bromoviridae

CMV infection in plants of Galium aparine was reported from New Zealand (Fletcher 2001). The virusinfected cleavers plants do not exhibit obvious symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

References Fletcher JD (2001) New hosts of Alfalfa mosaic virus, Cucumber mosaic virus, Potato virus Y, Soybean dwarf virus, and Tomato spotted wilt virus in New Zealand. N Z J Crop Hortic Sci 29:213–217

Gamochaeta spp. Family: Asteraceae

Weed host

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

Gardenia jasminoides (Cape jasmine)

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TSWV infection in plants of Gamochaeta spp. was reported from Georgia (USA) (Mullis et al. 2009). The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. For more details of TSWV, refer to Solanum lycopersicum.

References Mullis S, Bertrand PF, Brown SL, Csinos A, Diaz-Perez JC, Gitaitis R, Hickman LL, Johnson A, LaHue SS, Martinez N, McPherson RM, Nischwitz C, Reay-Jones F, Riley D, Sherwood J, Stevenson K, Wells L (2009) Tospoviruses in Solanaceae and other crops in the coastal plain of Georgia. University of Georgia, College of Agricultural and Environmental Sciences, Bulletin 1354, pp 51

G Gardenia jasminoides (Cape jasmine) Family: Rubiaceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

Infection of INSV was detected in plants of Gardenia jasminoides in New Zealand (Elliott et al. 2009). The virus-infected cape jasmine plants exhibit symptoms of mottling/mosaic, distortion, chlorosis, and stunting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Gardenia jasminoides was reported from Pennsylvania (USA) (Hausbeck et al. 1992). The virus-infected cape jasmine plants exhibit mottle-mosaic and necrosis symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Elliott DR, Lebas BSM, Ochoa-Corona FM, Tang J, Alexander BJR (2009) Investigation of Impatiens necrotic spot virus outbreaks in New Zealand. Aust Plant Pathol 38:490–495 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among greenhouse ornamentals in Pennsylvania. Plant Dis 76:795–800

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Gazania spp.

Gazania spp. Family: Asteraceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Gazania spp. was reported from Portugal (Louro 1996) and Iran (Shahraeen et al. 2002). The virus-infected gazania plants exhibit small necrotic spots, leaf yellowing, ringspots, necrotic vein-clearing, wilting, and dwarf symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Lettuce mosaic virus Taxonomic position Genus: Potyvirus

(LMV)

Family: Potyviridae

LMV infection in plants of Gazania spp. was reported from California, USA (Zerbini et al. 1993, 1997). The virus-infected gazania plants exhibit symptoms of yellow mottling or mosaic, chlorotic spots, and leaf distortion. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of LMV, refer to Lactuca sativa.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

Gazania spp. has been reported to be infected by TSWV in Iran, Italy, the Czech Republic, Greece, and Portugal (Louro 1996; Mertelík et al. 1996; Dellavalle et al. 1999; Chatzivassiliou et al. 2000; Ghotbi et al. 2005). The virus-infected gazania plants exhibit symptoms of necrotic spots and concentric ringspots, sometimes developing to necrotic patches. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Chatzivassiliou EK, Livieratos I, Jenser G, Katis NI (2000) Ornamental plants and thrips populations associated with Tomato spotted wilt virus in Greece. Phytoparasitica 28:257–264

Gentiana spp. (Gentian)

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Dellavalle G, Ciuffo M, Roggero P, Lisa V, Minuto A, Minuto G, Rapetti S (1999) Tospoviruses in Delphinium sp., gazania, marguerite, celery, Tragopogon porrifolius and Solanum rantonnetti in Liguria (Northern Italy). Inf Fitopatol 49:63–64 Ghotbi T, Sharaeen N, Winter S (2005) Occurrence of tospoviruses in ornamental and weed species in Markazi and Tehran provinces in Iran. Plant Dis 89:425–429 Louro D (1996) Detection and identification of tomato spotted wilt virus and Impatiens necrotic spot virus in Portugal. Acta Hortic 431:99–105 Mertelík J, Götsová B, Mokrá V (1996) Epidemiological aspects of Tomato spotted wilt virus infection in the Czech Republic. Acta Hortic 432:368–375 Shahraeen N, Ghotbi T, Mehraban AH (2002) Occurrence of Impatiens necrotic spot virus in ornamentals in Mahallat and Tehran Provinces in Iran. Plant Dis 86:694 Zerbini FM Jr, Koike ST, Gilbertson RL (1993) Gazania spp.: an alternate host of lettuce mosaic potyvirus (LMV) in the Salinas Valley. Phytopathology 83:1420 Zerbini FM, Koike ST, Gilbertson RL (1997) Gazania spp.: a new host of lettuce mosaic potyvirus, and a potential inoculum source for recent lettuce mosaic outbreaks in the Salinas Valley of California. Plant Dis 81:641–646

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Gentiana spp. (Gentian) Family: Gentianaceae

Ornamental

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

A potyvirus isolate causing gentian necrotic dwarf disease in Japan was initially stated to be an isolate of ClYVV and distinct from Bean yellow mosaic virus (BYMV) in several ways (Sasaya et al. 1997). However, further analysis showed this to be an isolate of BYMV (Uga et al. 2002). BYMV infection in plants of Gentian spp. was reported from Japan, Germany, the Netherlands, and the United Kingdom (Kaji et al. 1993; Uga et al. 2002; Verhoeven et al. 2002). The virus-infected gentian plants produce mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Broad bean wilt virus 2 Taxonomic position Genus: Fabavirus

(BBWV-2)

Family: Secoviridae

BBWV-2 infection in plants of Gentiana spp. was reported from Japan, the Netherlands, and Germany (Nagao et al. 1978; Yamashita 1990; Kobayashi et al. 1999; Verhoeven et al. 2002). The virus-infected gentian plants exhibit stunting and necrosis (from necrotic spots to whole-plant collapse) symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BBWV-2, refer to Vicia faba.

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Gentiana spp. (Gentian)

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Gentiana spp. was reported from Japan and New Zealand (Schmidt 1972; Nagao et al. 1978; Fletcher 1989; Yamashita 1990; Hikage et al. 2003; Sayama et al. 2006). The virusinfected gentian plants exhibit no symptoms or very mild symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sapinoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Gentian mosaic virus Taxonomic position Genus: Fabavirus

(GeMV)

Family: Secoviridae

Geographical distribution GeMV infection in plants of Gentiana scraba was reported from Japan (Kobayashi et al. 2005). Symptoms and host(s) The virus-infected gentian plants exhibit mosaic, necrotic spots, necrosis, leaf blight, and dwarfism symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. This virus is also mechanically sap-transmissible to nine plant species in six families (Solanaceae, Fabaceae, Chenopodiaeceae, Aizoaceae, Amaranthaceae, Gentianaceae). Chlorotic spots are produced on Chenopodium amaranticolor and C. quinoa. Virion properties and genome The virions are isometric and non-enveloped of two types but similar in size of 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is of bipartite, positive-strand RNA. RNA1 consists of 5,836 nt (AB084452) and RNA2 of 3,372 nt (AB084453). The virus particles consist of two coat proteins of 39.3 and 26.6 kDa (Ikegami and Sharma 2011; Kobayashi et al. 2005; Ferrer et al. 2007; Sanfacon et al. 2009; Sanfacon 2015; Thompson et al. 2017).

Gentian ovary ringspot virus Taxonomic position Genus: Goravirus

(GORV)

Family: Virgaviridae

Geographic distribution GORV was reported in plants of Gentiana triflora from Japan (Atsumi et al. 2015).

Gentiana spp. (Gentian)

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Symptoms and host(s) The virus-infected gentian plants cause unusual symptoms, ringspots that appear specifically on the outer surface of the ovarian wall after pollination.

Transmission There is no known vector for this virus. The virus was shown to be pollen transmitted. The virus could also be transmissible mechanically to petunia and tobacco.

Virion properties and genome The virions are rod-shaped and about 20 nm in width, but it was difficult to determine their length because of disintegration during purification. The genome has two single-stranded RNAs of 5519 nt (RNA 1; AB976029 = NC_024501) and 3869 nt (RNA 2; AB976030 = NC_024502) (Atsumi et al. 2015). RNA1 of GORV has two predicted ORFs, the first of which is a replication protein containing methyltransferase and helicase domains. The second is the RdRp; this is usually expressed as a readthrough product in the family Virgaviridae, and this may be the case with GORV although there is no direct experimental evidence. The first ORF of RNA2 is a coat protein, and subsequent ORFs encode the triple gene block proteins (involved in virus cell-to-cell movement) and a cysteine-rich protein (CRP) that was shown to act as suppressor of gene silencing (Atsumi et al. 2015; Adams et al. 2017).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Gentiana spp. was reported from the Netherlands, China, and Japan (Verhoeven et al. 2002; Nekoduka et al. 2005; Ding et al. 2011). The virus-infected gentian plants exhibit symptoms of chlorotic and necrotic spotting, necrotic mottle, and severe stunting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in field-grown plants of Gentiana spp. was reported from Japan (Kuroda et al. 2002; Uga 2005). The virus is transmitted by nematode vectors and is also transmissible by mechanical sapinoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

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Gentiana spp. (Gentian)

Tomato black ring virus Taxonomic position Genus: Nepovirus

(TBRV)

Family: Secoviridae

TBRV was detected in a single plant of Gentiana asclepiadea from Germany (Verhoeven et al. 2002). The virus-infected gentian plants exhibit chlorotic mottle and stunting symptoms. The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of TBRV, refer to Solanum lycopersicum.

References Adams MJ, Adkins S, Bragard C, Gilmer D, Li D, MacFarlane SA, Wong S-M, Melcher U, Ratti C, Ryu KH, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Virgaviridae. J Gen Virol 98:1999–2000 Atsumi G, Tomita R, Yamashita T, Sekine K-T (2015) A novel virus transmitted through pollination causes ring-spot disease on gentian (Gentiana triflora) ovaries. J Gen Virol 96:431–439 Ding M, Yin Y, Fang Q, Li S, Zhang Z (2011) First report of impatiens necrotic spot virus in Gentiana macrophylla in China. Plant Dis 95:357 Ferrer RM, Luis-Arteaga M, Guerri J, Moreno P, Rubio L (2007) Detection and identification of species of the genus Fabavirus by RT–PCR with a single pair of primers. J Virol Methods 144:156–160 Fletcher JD (1989) Additional hosts of Alfalfa mosaic virus, Cucumber mosaic virus and Tobacco mosaic virus in New Zealand. N Z J Crop Hortic Sci 17:361–362 Hikage T, Yoshiike T, Kominato M, Atarashi H, Sayama H (2003) Six-year protective effect of attenuated Cucumber mosaic virus strains in gentian plants. Jpn J Phytopathol 69:320–321 Ikegami M, Sharma P (2011) Fabavirus. Comoviridae. In: Tidona C, Darai G (eds) The Springer Index of Viruses. Springer, New York, NY. https://doi.org/10.1007/978-0-387-95919-1 Kaji K, Sonoda T, Fujisawa I (1993) Gentian necrotic dwarf disease caused by Bean yellow mosaic virus N strain. Ann Phytopathol Soc Jpn 59:57 Kobayashi YO, Nakano M, Kashiwazaki S, Naito T, Mikoshiba Y, Shiota A, Kameya-Iwaki M, Honda Y (1999) Sequence analysis of RNA-2 of different isolates of broad bean wilt virus confirms the existence of two distinct species. Arch Virol 144:1429–1438 Kobayashi YO, Kobayashi A, Hagiwara K, Ugah Mikoshiba Y, Naito T, Honda Y, Omura T (2005) Gentian mosaic virus: A new species in the genus Fabavirus. Phytopathology 95:192–197 Kuroda T, Urushibara S, Takeda I, Nakatani F, Suzuki K (2002) Multiplex reverse transcription polymerase chain reaction for simultaneous detection of viruses in gentian. J Gen Plant Pathol 68:169–172 Nagao N, Torigoe H, Wakimoto S (1978) Cucumber mosaic virus and broad bean wilt virus isolated from gentian. Proc Assoc Plant Protect Kyushu 24:180 Nekoduka S, Hamada H, Katsube K (2005) Necrotic mottle disease of gentian caused by Impatiens necrotic spot virus (INSV) (in Japanese with English summary). Jpn J Phytopathol 71:183–184 Sanfacon H (2015) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. In: Encyclopedia of Life Sciences. Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000764.pub3. http:// www.els.net Sanfacon H, Wellink J, Le Gall O, Karasev A, van der Vlugt R, Wetzel T (2009) Secoviridae: a proposed family of plant viruses within the order Picornavirales that combines the families Sequiviridae and Comoviridae, the unassigned genera Cheravirus and Sadwavirus, and the proposed genus Torradovirus. Arch Virol 154(5):899–907 Sasaya T, Kaji K, Fujisawa I, Koganezawa H, Teraoka T, Natsuaki T, Okuda S (1997) Characterization and nucleotide sequence of the 30 terminal region of Clover yellow vein virus isolated from Gentian (Gentiana spp.). Jpn J Phytopathol 63:328–333 Sayama H, Kominato M, Atarashi H, Takayanagi N, Yamada M, Hikage T, Yoshiike T (2006) Control of cucumber mosaic virus (CMV) in gentian by satellite RNA containing attenuated CMV strains. Acta Hortic 722:147–153 Schmidt HE (1972) Mottled crinkle of stemless gentian (Gentiana alpine Vill.) induced by Cucumber mosaic virus. Arch Gartenbau 20:575–579 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Secoviridae. J Gen Virol 98:529–531

Gerbera spp. (Gerbera)

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Uga H (2005) Study on viral diseases of dwarf gentian plants and their control using attenuated isolate. Bulletin of the Saitama Prefectural Agriculture and Forestry Research Center, pp 1–85 Uga H, Kobayashi YO, Hagiwara K, Honda Y, Omura T, Sasaya T (2002) Molecular characterization of Bean yellow mosaic virus from vegetatively propagated dwarf Gentiana plants. J Gen Plant Pathol 68:378–381 Verhoeven JTHJ, Ding YM, Lesemann D-E, van der Vlugt RAA, Roenhorst JW (2002) Virus infections in Gentiana species. Acta Hortic 568:61–67 Yamashita K (1990) Mosaic diseases occurred on gentian and prairie gentian in Aomori Prefecture. Ann Rep Soc Plant Protect N Jpn 41:80–82

Geranium carolinianum (Carolina cranesbill) Family: Geraniaceae

Weed host

G Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Geranium carolinianum was reported from the USA (Groves et al. 2002; Mullis et al. 2009). The virus-infected Carolina cranesbill plants exhibit yellow spots or ringspot on the leaves. The virus is transmitted by the thrips vector, Frankliniella fusca, in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Groves RL, Walgenbach JF, Mayor JW, Kennedy GG (2002) The role of weed hosts and tobacco thrips, Frankliniella fusca, in the epidemiology of Tomato spotted wilt virus. Plant Dis 86:573–582 Mullis S, Bertrand PF, Brown SL, Csinos A, Diaz-Perez JC, Gitaitis R, Hickman LL, Johnson A, LaHue SS, Martinez N, McPherson RM, Nischwitz C, Reay-Jones F, Riley D, Sherwood J, Stevenson K, Wells L (2009) Tospoviruses in Solanaceae and other crops in the coastal plain of Georgia. University of Georgia, College of Agricultural and Environmental Sciences, Bulletin 1354, pp 51

Gerbera spp. (Gerbera) Family: Asteraceae

Ornamental

Chrysanthemum stem necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(CSNV)

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Gerbera spp. (Gerbera)

CSNV infection in plants of Gerbera spp. was reported from Slovenia (Boben et al. 2007). The infected plants show symptoms typical of tospovirus infection: slight yellowing of leaves and necrosis. The virus is efficiently transmitted by Frankliniella occidentalis and F. schultzeri in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of CSNV, refer to Chrysanthemum spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Gerbera spp. was reported from Australia, India, and Cuba (Finlay 1975; Verma et al. 2004; González Arias and del Loreto Reyes 2013; Gautam et al. 2014). The virus-infected gerbera plants exhibit severe chlorotic mosaic and flower deformation symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Gerbera symptomless virus

(GeSLV)

Synonyms Gerbera latent virus Taxonomic position GeSLV is a tentative member of the family Rhabdoviridae. Geographical distribution GeSLV was first reported in plants of Gerbera spp. from Japan by Chang et al. (1976). Symptoms and host(s) The virus-infected gerbera plants exhibit symptomless infections. Virion properties and genome The virions are c.60–70  150–300 bacilliform or bullet-shaped. No specific genome information is available but anticipated to be a negative-stranded RNA of >12 kb.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Gerbera spp. has been reported in the USA, Portugal (Louro 1996), and New Zealand (Elliott et al. 2009). The virus-infected gerbera plants exhibit symptoms which may begin as light green mottle, followed by necrotic ring and line patterns. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Gerbera spp. (Gerbera)

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Tobacco mosaic virus

(TMV)

Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

TMV infection in plants of Gerbera jamesonii was reported from Turkey (Yorganici and Karaca 1974; Sutic et al. 1999). The virus-infected gerbera plants exhibit foliar mottle or mosaic symptoms. There is no known vector for this virus. The virus is mechanically sap-transmissible and is also transmissible by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco rattle virus

(TRV)

G

Taxonomic position Genus: Tobravirus

Family: Virgaviridae

TRV was isolated from plants of Gerbera jamesonii in the USA (Stouffer 1965) and the Netherlands (Hakkaart 1968). The virus-infected gerbera plants exhibit black or yellow ringspots on the leaves. The virus is transmitted by nematode vectors and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Gerbera spp. was reported from Iran (Ghotbi and Shahraeen 2009). The virus-infected gerbera plants exhibit symptoms of general yellowing and leaf discoloration. The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato black ring virus Taxonomic position Genus: Nepovirus

(TBRV)

Family: Secoviridae

TBRV infection in plants of Gerbera jamesonii was reported from China (Zhang et al. 2009). The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of TBRV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

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Gesneria spp.

TSWV infection in plants of Gerbera jamesonii was reported from Venezuela, Pennsylvania (USA), Portugal, Italy, and Serbia (Hausbeck et al. 1992; Louro 1996; Spano et al. 2011; Stankovic et al. 2011; Marys et al. 2014). The virus-infected gerbera plants show concentric rings, irregular chlorotic blotches, and deformation on leaves. The virus is transmitted by a thrips vector Frankliniella occidentalis in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of TSWV, refer to Solanum lycopersicum.

References Boben J, Mehle N, Pirc M, Mavrič Pleško I, Ravnikar M (2007) New molecular diagnostic methods for detection of Chrysanthemum stem necrosis virus (CSNV). Acta Biol Slov 50:41–51 Chang MU, Doi Y, Yora K (1976) Ann Phytopathol Soc Jpn 42:383 Elliott DR, Lebas BSM, Ochoa-Corona FM, Tang J, Alexander BJR (2009) Investigation of Impatiens necrotic spot virus outbreaks in New Zealand. Australas Plant Pathol 38:490–495 Finlay JR (1975) Cucumber mosaic virus in gerbera. Aust Plant Pathol 4:14 Gautam KK, Raj R, Kumar S, Raj SK, Roy RK, Katiyar R (2014) Complete sequence of RNA3 of Cucumber mosaic virus isolates infecting Gerbera jamesonii suggests its grouping under IB subgroup. Virus Dis 25:398–401 Ghotbi T, Shahraeen N (2009) Natural incidence and infectivity level of three nepoviruses in ornamental crops in Iran. J Plant Breed Crop Sci 1:39–44 González Arias G, del Loreto Reyes M (2013) Genera and species of virus present in ornamental plants and cut flowers in the provinces of La Habana and Cienfuegos, Cuba. Fitosanidad 17:103–105 Hakkaart FA (1968) A virus disease of Gerbera jamesonii. Neth J Plant Pathol 74:28–29 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among greenhouse ornamentals in Pennsylvania. Plant Dis 76:795–800 Louro D (1996) Detection and identification of tomato spotted wilt virus and Impatiens necrotic spot virus in Portugal. Acta Hortic 431:99–105 Marys E, Mejias A, Rodriguez-Roman E, Avilan D (2014) The first report of Tomato spotted wilt virus on Gerbera and Chrysanthemun in Venezuela. Plant Dis 98:1161 Spano R, Mascia T, De Lucia B, Torchetti EM, Rubino L, Gallitelli D (2011) Disease note: first report of a resistancebreaking strain of Tomato spotted wilt virus from Gerbera jamesonii in Apulia, southern Italy. J Plant Pathol 93(4 supplement):S4.63 Stankovic I, Bulajic A, Vucurovic A, Ristic D, Jovic J, Krstic B (2011) First report of Tomato spotted wilt virus on Gerbera hybrida in Serbia. Plant Dis 95:226 Stouffer RF (1965) Isolation of tobacco rattle virus from Transvaal daisy, Gerbera jamesonii. Phytopathology 55:501 Sutic DD, Ford RE, Tosic MT (1999) Tobacco mosaic tobamovirus (TMV). In: Handbook of plant virus diseases. CRC Press, Florida, USA Verma N, Singh AK, Singh L, Kulshreshtha S, Raikhy G, Hallan V, Ram R, Zaidi AA (2004) Occurrence of Cucumber mosaic virus in Gerbera jamesonii in India. Plant Dis 88:1161 Yorganici U, Karaca I (1974) Tobacco mosaic virus on Gerbera jamesonii in Turkey. J Turk Phytopathol 3:116–123 Zhang LL, Yang CY, Yu C, Yuan WH, Ma Q (2009) Rapid detection of 3 viruses in Gerbera jamesonii Bolus and its virusfree by tissue culture. Acta Phytophylacica Sin 36:239–245

Gesneria spp. Family: Gesneriaceae

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

Ornamental

(TMV)

Family: Virgaviridae

Gladiolus spp.

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TMV infection in plants of Gesneria spp. was reported from Florida (USA) (Smith 1957a, b; Zettler and Nagel 1983). The symptoms on the foliage of infected gesneria plants were inconspicuous. There is no known vector for this virus. The virus is mechanically sap-transmissible and is also transmissible by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

References Smith KM (1957a) A textbook of plant virus diseases. Little, Brown & Co., Boston 652 pp Smith KM (1957b) A text book of plant virus diseases, 2nd edn. Churchill, London 572 pp Zettler FW, Nagel J (1983) Infection of cultivated gesneriads by two strains of Tobacco mosaic virus. Plant Dis 67:1123–1125

Gibasis geniculata (Tahitian bridal veil) Family: Commelinaceae

Ornamental

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Gibasis geniculata was reported from Virginia (USA) (Hunst and Tolin 1982). The virus-infected Tahitian bridal veil plants exhibit severe stunting and leaf distortion symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

References Hunst PL, Tolin SA (1982) Bean yellow mosaic virus isolated from Gibasis geniculata. Plant Dis 66:955–958

Gladiolus spp. Family: Iridaceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

Family: Secoviridae

ArMV infection in plants of Gladiolus spp. has been reported from India, Italy, Lithuania, and Iran (Bellardi and Marani 1984; Katoch et al. 2003b; Ghotbi and Shahraeen 2005, 2009; Samuitiene et al. 2008). Two types of isolates were reported from Italy; one type was found in a single infection

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Gladiolus spp.

associated with mild mosaic, whereas the other was always found in a mixed infection with BYMV in plants with marked leaf symptoms (Bellardi et al. 1986). Light flower break was observed in some plants with colored, but not white, flowers. This virus is transmitted by the nematode vectors (Xiphinema spp.) in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Bean common mosaic virus Taxonomic position Genus: Potyvirus

(BCMV)

Family: Potyviridae

BCMV infection in plants of Gladiolus spp. was reported from Oregon, USA (Snow 1956). The virusinfected gladiolus plants exhibit symptoms of foliage strongly mottled with dark green rectangular block patterns formed by bleached, semi-transparent, yellowish squares of tissue. The normally pink flower was weakly broken by yellowish clear patches of tissue. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BCMV, refer to Phaseolus vulgaris.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Gladiolus spp. was distributed worldwide (Zettler and Abo El-Nil 1977; Stein et al. 1979, 1986, 1994; Kaniran and Izadpanah 1982; Srivastava et al. 1983; Hseu et al. 1987; Raizada et al. 1989; Park and Kim 1998; Navalinskiene and Samuitiene 2001; Katoch et al. 2002; Hammond and Hammond 2003; Arneodo et al. 2005; Dubey et al. 2009; Duraisamy and Pokorny 2009; Dorrigiv et al. 2013; Kaur et al. 2015). Some virus-infected gladiolus plants exhibit flower colorbreaking symptoms, whereas others exhibit streaks, mosaics, mottling, white-breaking symptoms on leaves, and leaf malformation, while some gladiolus varieties were symptomless (Magie and Poe 1972; Vunsh et al. 1991; Selvarajan and Gupta 1996). The virus is transmitted by aphids including Acyrthosiphon pisum, Macrosiphum euphorbiae, Myzus persicae, and Aphis fabae, in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. The use of corms from the infected plants is a major cause of virus spread. For more details of BYMV, refer to Phaseolus vulgaris.

Clover yellow vein virus Taxonomic position Genus: Potyvirus

(ClYVV)

Family: Potyviridae

ClYVV was detected in plants of Gladiolus spp. in Korea (Park and Kim 1998). The virus-infected gladiolus plants show flower color breaking, leaf mosaic, necrotic fleck, and dwarfing or lack of visible symptoms. The virus is transmitted by various aphids in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ClYVV, refer to Trifolium spp.

Gladiolus spp.

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Cucumber mosaic virus

(CMV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMVoccurs wherever Gladiolus spp. are cultivated (Brierley 1952; McWhorter 1957; Stein et al. 1979; Stein 1995; Park et al. 1998; Navalinskiene and Samuitiene 2001; De la Torre-Almaraz et al. 2002; Raj et al. 2002; Arneodo et al. 2005; Samuitiene and Navalinskiene 2008; Dubey et al. 2008, 2010; Duraisamy and Pokorny 2009; Rishi 2009; Dorrigiv et al. 2013). The virus-infected gladiolus plants exhibit symptoms of small chlorotic squares between parallel leaf veins. The spots may be gray, yellow, brown, or reddish. When the symptoms are severe, the plant may have an overall yellow appearance. To detect mild symptoms, a leaf must be held up to the light. Stunted plants are common, and flower discoloration may be accompanied by crinkling, shrinking, or flower breaking/deformations. The corms from infected plants of some varieties may be warty when harvested. The virus is transmitted to gladiolus plants by the feeding of aphids, such as Myzus persicae, Aphis craccivora, etc., in a non-persistent manner. The virus is transmissible mechanically by sapinoculation, and also by contaminated tools. For more details of CMV, refer to Cucumis sativus.

Cycas necrotic stunt virus Taxonomic position Genus: Nepovirus

(CNSV)

Family: Secoviridae

CNSV has been reported from plants of Gladiolus spp. only in Japan (Fukumoto et al. 1982; Hanada et al. 2006). The virus is mechanically sap-transmissible. For more details of CNSV, refer to Cycas spp.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Gladiolus spp. was reported from Portugal and Iran (Louro 1996; Ghotbi et al. 2005; Ghotbi 2013). The virus-infected gladiolus plants exhibit symptoms of necrotic leaf spot, chlorosis, and stunting. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Ornithogalum mosaic virus Taxonomic position Genus: Potyvirus

(OrMV)

Family: Potyviridae

OrMV occurs in plants of Gladiolus spp. in the Netherlands, South Africa, the USA, and India (Kaur et al. 2011). The virus-infected gladiolus plants show symptoms of mosaic, chlorotic spots on leaves,

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Gladiolus spp.

and floral deformations. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of OrMV, refer to Ornithogalum spp.

Tobacco necrosis virus

(TNV)

Taxonomic position TNV is a tentative member of the genus Necrovirus and family Tombusviridae. TNV infection in plants of Gladiolus spp. was reported from Russia and Lithuania (Navalinskiene and Samuitiene 2010). The virus-infected gladiolus plants show light green stripes and spots on leaves and sometimes brown necrotic pinpoint spots. Later in the season, the necrotic spots expanded and coalesced, causing severe symptoms. It is not clear whether the isolates from gladiolus are of TNV-A (Alphanecrovirus) or TNV-D (Betanecrovirus). The virus is transmitted by the zoospores of the fungus Olpidium brassicae and is also transmissible by mechanical sap-inoculation. For more details of TNV, refer to Nicotiana tabacum.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in plants of Gladiolus spp. was reported from the Netherlands, India, Israel, Egypt, the Czech Republic, and Poland (Asjes 1994; Park et al. 1998; Yamaji et al. 1998; Navalinskiene and Samuitiene 2001; Katoch et al. 2004; Duraisamy and Pokorny 2009). The virus-infected gladiolus plants show notched leaf blade margins. The virus is transmitted by nematode vectors. The virus is mechanically sap-transmissible, and inoculated Nicotiana tabacum produces symptoms of necrotic local lesions followed by systemic mottling symptoms. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Gladiolus spp. was reported from the USA, India, and Lithuania (Brierley 1952; Fukumoto et al. 1982; Kaniran and Izadpanah 1982; Katoch et al. 2003a, b; Navalinskiene and Samuitiene 2010). The virus-infected gladiolus plants develop necrotic or chlorotic ringspot patterns on leaves. The flowers are not affected. Mild strains of the virus may cause mild chlorotic mottling in gladiolus. The virus is transmitted through the nematode vector, Xiphinema americanum, in a nonpersistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Gladiolus spp.

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Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Gladiolus spp. was reported from Italy and Iran (Bellardi et al. 1987; Ghotbi 2006). The virus-infected gladiolus plants exhibit stunting of flower heads and wavelike necrotic rings or line patterns in the leaves. The virus is transmitted by the thrips vectors. The virus present in/on pollen entering the host through injuries caused by thrips while feeding. The virus is also mechanically sap-transmissible, but is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato aspermy virus

(TAV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

TAV infection in plants of Gladiolus spp. was reported from India (Raj et al. 2011). The virus-infected gladiolus plants exhibit symptoms of severe mosaic, with necrotic dots along the midrib of leaves. Severely infected plants bore very few and small florets with color-breaking symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of TAV, refer to Solanum lycopersicum.

Tomato black ring virus Taxonomic position Genus: Nepovirus

(TBRV)

Family: Secoviridae

TBRV infection in plants of Gladiolus spp. was reported from Poland, causing symptoms on both leaves and flowers (Kaminska 1978), but later reported from Italy in symptomless plants (Bellardi and Pisi 1985). The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of TBRV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Gladiolus spp. was reported from the USA and Iran (Bozarth and Corbett 1957; Kaniran and Izadpanah 1982; Loebenstein et al. 1995; Ghotbi et al. 2005; Ghotbi and Shahraeen 2009). The virus-infected gladiolus plants exhibit symptoms of stunting and have short flower spikes. Leaves on affected plants are usually smaller, stiffer, and more erect. The virus is transmitted by the nematode vectors Xiphinema and Dorylamidae spp. in a non-persistent manner, and is also

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Gladiolus spp.

transmissible by mechanical sap-inoculation. The virus is transmitted by seed and pollen. For more details of ToRSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Gladiolus spp. was reported from Australia, Portugal, and the USA (Lee et al. 1979; Louro 1996; Katoch et al. 2003b; Parrella et al. 2003). The virus-infected gladiolus plants show chlorotic degeneration, occurring at a high incidence in one season, though previously present in other crops without obvious disease in gladiolus. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is not transmissible by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Arneodo JD, de Breuil S, Lenardon SL, Conci VC, Conci LR (2005) Detection of Bean yellow mosaic virus and Cucumber mosaic virus infecting gladiolus in Argentina. Agriscientia 22:87–89 Asjes CJ (1994) Occurrence of Tobacco rattle virus in ornamental bulbous crops in the Netherlands. Acta Hortic 377:349–356 Bellardi MG, Marani F (1984) Nepoviruses isolated from gladiolus in Italy. Acta Hortic 164:297–308 Bellardi MG, Pisi A (1985) Identification of three isolates of tomato black ring virus (TBRV) from gladiolus. Riv Ortoflorofrutticoltura Ital 69:299–309 Bellardi MG, Canova A, Gelli C (1986) Comparative studies on gladiolus isolates of Arabis mosaic virus (ArMV). Phytopathol Mediterr 25:85–91 Bellardi MG, Vicchi V, Gelli C (1987) Stunting of gladiolus flower spike associated with tobacco streak virus (TSV). Phytopathol Mediterr 26:73–80 Bozarth RF, Corbett MK (1957) Tomato ringspot virus associated with stunt or stub disease of gladiolus in Florida. Plant Dis Reptr 42:217–221 Brierley P (1952) Evidence on the significance of cucumber mosaic and tobacco ringspot viruses in gladiolus. USDA Plant Dis Reptr 36(2):48–50 De la Torre-Almaraz R, Cruz Monsalvo-Reyes R, Salazar-Segura M, Valverde RA (2002) Detection and partial characterization of a yellow variant of Cucumber mosaic virus in gladiola (Gladiolus grandiflorus Hort.) in Mexico. Rev Chapingo Ser Hortic 8(1):83–93 Dorrigiv R, Jafarpour B, Rastegar MF (2013) Detection of some virus pathogens of gladiolus in Iran. Int J Agric Crop Sci 5:1653–1658 Dubey VK, Aminuddin, Singh VP (2008) First report of a subgroup IA Cucumber mosaic virus isolate from gladiolus in India. Australas Plant Dis 3:35–37 Dubey VK, Aminuddin A, Singh VP (2009) First report of Bean yellow mosaic virus in gladiolus in India. J Plant Pathol 91(4):787–791 Dubey VK, Aminuddin, Singh VP (2010) Molecular characterization of Cucumber mosaic virus infecting Gladiolus, revealing its phylogeny distinct from the Indian isolate and alike the Fny strain of CMV. Virus Genes 41:126–134 Duraisamy GA, Pokorny R (2009) Survey of virus pathogens in gladiolus, iris, and tulip in the Czech Republic. Acta Univ Agric Silviculturae Mendelianae Brunensis 57:79–86 Fukumoto F, Ito Y, Tochihara H (1982) Viruses isolated from Gladiolus in Japan. Ann Phytopathol Soc Jpn 48:68–71 Ghotbi T (2006) First report on incidence of Tobacco streak virus (TSV) on ornamental plants in Iran. Iran J Plant Pathol 42:159–160 Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iran J Plant Pathol 49:41 Ghotbi T, Shahraeen N (2005) First report on incidence of Arabis mosaic virus (ArMV, Nepovirus) on ornamental plants in Iran. Iran J Plant Pathol 41(2):305–306

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Ghotbi T, Shahraeen N (2009) Natural incidence and infectivity level of three nepoviruses in ornamental crops in Iran. J Plant Breed Crop Sci 1(3):39–44 Ghotbi T, Sharaeen N, Winter S (2005) Occurrence of tospoviruses in ornamental and weed species in Markazi and Tehran provinces in Iran. Plant Dis 89:425–429 Hammond J, Hammond RW (2003) The complete nucleotide sequence of isolate BYMV-GDD of Bean yellow mosaic virus, and comparison to other potyviruses. Arch Virol 148:2461–2470 Hanada K, Fukumoto F, Kusunoki M, Kameya-Iwaki M, Tanaka Y, Iwanami T (2006) Cycas necrotic stunt virus isolated from gladiolus plants in Japan. J Gen Plant Pathol 72:383–386 Hseu SH, Huang CH, Chang CA (1987) Bean yellow mosaic virus from gladiolus: identification, purification and its serological properties. J Agric Res China 36:94 Kaminska M (1978) Some properties of isolates of tomato black ring virus. Zesz Probl Postep Nauk Rol 214:109–117 Kaniran R, Izadpanah K (1982) Isolation and identification of bean yellow mosaic and tobacco ringspot viruses from gladiolus in Shiraz. Iran J Plant Pathol 17:1–10 Katoch M, Ram R, Zaidi AA, Garg ID (2002) Status of Bean yellow mosaic virus on Gladiolus. Crop Protect 21:861–865 Katoch M, Ram R, Zaidi AA (2003a) First report of Tobacco ringspot virus in Gladiolus in India. Plant Pathol 52:789 Katoch M, Abdin MZ, Ram R, Zaidi AA (2003b) An overview of diagnostics for viruses infecting gladiolus. Crop Protect 22:153–156 Katoch M, Abdin MZ, Zaidi AA (2004) First report of Tobacco rattle virus occurring in gladiolus in India. Plant Pathol 53:236 Kaur C, Raj SK, Snehi SK, Goel AK, Roy RK (2011) Natural occurrence of Ornithogalum mosaic virus newly reported on gladiolus in India. New Dis Rep 24:2 Kaur C, Kumar S, Raj SK, Chauhan PS, Sharma N (2015) Characterization of a new isolate of Bean yellow mosaic virus group-IV associated with mosaic disease of gladiolus in India. J Plant Pathol Microbiol 6(10):309 Lee TC, Francki RIB, Hatta T (1979) A serious disease of gladiolus in Australia caused by tomato spotted wilt virus. Plant Dis Reptr 63:343–348 Loebenstein G, Lawson RH, Brunt AA (1995) Virus and virus-like diseases of bulb and flowering crops. Wiley, Chichester, p 543 Louro D (1996) Detection and identification of tomato spotted wilt virus and Impatiens necrotic spot virus in Portugal. Acta Hortic 431:99–105 Magie RO, Poe SL (1972) Disease and pest associates of bulbs and plants. In: Koenig R, Crowley W (eds) The world of gladiolus. North American Gladiolus Council, Edgewood Press, Edgewood, pp 155–167 McWhorter FP (1957) A localized occurrence of Cucumber mosaic virus in Gladiolus. Plant Dis Reptr 41:141–143 Navalinskiene M, Samuitiene M (2001) Viral diseases of flower plants 15. Identification of viruses affecting gladiolus (Gladiolus L.). Biologija 1:31–35 Navalinskiene M, Samuitiene M (2010) Identification of viral and phytoplasmal diseases affecting gladioli (Gladiolus L.). Acta Biol Univ Daugavp Suppl 2:137–144 Park IS, Kim KW (1998) The viruses in Gladiolus hybridus cultivated in Korea 1. Bean yellow mosaic virus and clover yellow vein virus. Korean J Plant Pathol 14:74–82 Park I, Kim K, Kyun H, Chang M (1998) The viruses in Gladiolus hybridus cultivated in Korea. 2. Broad bean wilt virus, cucumber mosaic virus and tobacco rattle virus. Korean J Plant Pathol 14:83–91 Parrella G, Gognalons P, Gebre-Selassi K, Vovlas C, Marchoux G (2003) An update of the host range of tomato spotted wilt virus. J Plant Pathol 85(4, special issue):227–264 Raizada RK, Zaidi AA, Srivastava KM, Shreni VCD, Singh BP (1989) Detection of Bean yellow mosaic virus in different parts of gladiolus. Indian J Plant Pathol 7:91–96 Raj SK, Srivastava A, Chandra G, Singh BP (2002) Characterization of Cucumber mosaic virus isolate infecting gladiolus cultivars and comparative evaluation of serological and molecular methods for sensitive diagnosis. Curr Sci 83:1132–1137 Raj SK, Kumar S, Verma DK, Snehi SK (2011) First report on molecular detection and identification of Tomato aspermy virus naturally occurring on gladiolus in India. Phytoparasitica 39:303–307 Rishi N (2009) Significant plant virus diseases in India and a glimpse of modern disease management technology. J Gen Plant Pathol 75:1–18 Samuitiene M, Navalinskiene M (2008) Occurrence of Cucumber mosaic cucumovirus on ornamental plants in Lithuania. Zemdirbyste-Agriculture 95:135–143 Samuitiene M, Navalinskiene M, Jackeviciene E (2008) Arabis mosaic virus on ornamental plants. Biologia 54:264–268 Selvarajan R, Gupta MD (1996) Characterization of a virus causing mosaic on gladiolus. Int J Trop Plant Dis 14:217–222 Snow GF (1956) Ringspot and bean mosaic virus from gladiolus. PhD thesis, Oregon State College, Corvallis, 80 p Srivastava KM, Raizada RK, Singh BP (1983) Bean yellow mosaic virus on Gladiolus psittacinus in India. Indian J Plant Pathol 1:83–88 Stein A (1995) Gladiolus. In: Loebenstein G, Lawson RH, Brunt AA (eds) Virus and virus-like diseases of bulb and flowering crops. Wiley, Chichester, pp 281–292, 543 pp

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Globularia nudicaulis (Globe daisy)

Stein A, Loebenstein G, Koenig R (1979) Detection of Cucumber mosaic virus and Bean yellow mosaic virus in gladiolus by enzyme linked immunosorbent assay (ELISA). Plant Dis Reptr 63:185–188 Stein A, Salomon R, Cohen J, Loebenstein G (1986) Detection and characterisation of Bean yellow mosaic virus in corms of Gladiolus grandiflorus. Ann Appl Biol 109:147–154 Stein A, Rosner A, Hammond J (1994) Detection of bean yellow mosaic virus in gladioli corms. Acta Hortic 377:209–220 Vunsh R, Rosner A, Stein A (1991) Detection of bean yellow mosaic virus in gladioli corms by the polymerase chain reaction. Ann Appl Biol 119:289–294 Yamaji Y, Horikoshi K, Yamashita H, Matsumoto T (1998) Detection of tobacco rattle virus on gladiolus by RT-PCR. Res Bull Plant Protect Serv Jpn 0(34):107–111 Zettler FW, Abo El-Nil MM (1977) Bean yellow mosaic virus infections on gladiolus in Florida. Plant Dis Reptr 61:243–247

Globularia nudicaulis (Globe daisy) Family: Plantaginaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Weed host

(CMV)

Family: Bromoviridae

CMV infection in plants of Globularia nudicaulis was reported from Italy (Davino et al. 2006). The virus-infected globe daisy plants exhibit symptoms of yellow mosaic and variegation on malformed leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

References Davino S, Cugnata S, Bellardi MG (2006) Globularia nudicaulis, a new host for Cucumber mosaic virus. Plant Pathol 55:568

Gloriosa spp. (Gloriosa rothschildiana; G. superba) (Gloriosa lily) Family: Colchicaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Gloriosa rothschildiana was reported from Japan (Araki et al. 1985). The virus-infected gloriosa lily plants show mottling symptoms. The virus is transmitted by a number of

Gloriosa spp. (Gloriosa rothschildiana; G. superba) (Gloriosa lily)

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aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Gloriosa stripe mosaic virus

(GSMV)

Synonyms Gloriosa lily mosaic virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution GSMV infection in plants of Gloriosa spp. was reported from Germany (Koenig and Lesemann 1974), Japan (Araki et al. 1985), the Netherlands (Pham et al. 2011), and the USA (Mollov et al. 2017). Symptoms and host(s) The virus-infected gloriosa lily plants exhibit stripe mosaic or mottle symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is transmissible to a healthy Gloriosa by mechanical inoculation. Virion properties and genome The virions are non-enveloped, flexuous filaments, 760 nm long and 13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9455 nt (EF427894). Partial sequences of German, Dutch, and US isolates are also available (EU042761, EU042762; KT602853-KT602855). The partial sequence of the Dutch isolate (EU042761) is about 81% identical to a Potyvirus affecting G. superba in Taiwan, described as Glory lily mosaic virus (EU250360), suggesting synonymy (Pham et al. 2011; Wylie et al. 2017).

References Araki M, Yamashita S, Doi Y, Yora K (1985) Three viruses from Gloriosa (Gloriosa rothschildiana O’brien) Gloriosa fleck virus, Gloriosa stripe mosaic virus and Cucumber mosaic virus. Jpn J Phytopathol 51:632–636 Koenig R, Lesemann D (1974) A potyvirus from Gloriosa rothschildiana. Phytopathol Z 80:136–142 Mollov D, Grinstead S, Tahir MN, Reinsel MD, Hammond J (2017) First report of multiple isolates of Gloriosa stripe mosaic virus in Gloriosa superba in the United States. Plant Dis 101:1070 Pham KTK, de Kock MJD, Lemmers MEC, Derks AFLM (2011) Molecular identification of potyviruses infecting bulbous ornamentals by analysis of coat protein (CP) sequences. Acta Hortic 901:167–172 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354

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Gloxinia spp.

Gloxinia spp. Family: Gesneriaceae

Ornamental

Capsicum chlorosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(CaCV)

Family: Tospoviridae

CaCV was originally described in the USA as isolate HT-1 and separated from a mixed infection with INSV (Hsu et al. 2000). A Tospovirus from gloxinia was later identified as Capsicum chlorosis virus (Chen et al. 2006, 2012). The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of CaCV, refer to Capsicum annuum.

Columnea latent viroid Taxonomic position Genus: Pospiviroid

(CLVd)

Family: Pospiviroidae

CLVd was detected in plants of Gloxinia spp. from Denmark, South America, and Bolivia (Nielsen and Nicolaisen 2010). The viroid was identified in G. gymnostoma, G. nematanthodes, and G. purpurascens and was symptomless in these ornamental species. The viroid is mechanically saptransmissible and also spreads easily via contaminated cutting tools while handing or through direct plant-to-plant contact. For more details of CLVd, refer to Columnea spp.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Gloxinia spp. was reported from the USA (Daughtrey et al. 1997). The virus-infected gloxinia plants exhibit chlorotic and brown ringspots, line patterns, mosaic, veinal necrosis, distortion of young growth, stunting, and wilting. The virus is transmitted by the thrips vector, Frankliniella occidentalis, in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV was detected in the USA in cultivated Gloxinia spp., but not in any of five specimens of wild origin in Central or South America (Zettler and Nagel 1983). The symptoms on the foliage of infected gloxinia

Glycine clandestina (Twining glycine)

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plants were inconspicuous. There is no known vector for this virus. The virus is mechanically saptransmissible and is also transmissible by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Gloxinia spp. was reported from Europe and the USA (Jones and Moyer 1986; Brown 1988; Bertaccini and Bellardi 1990; Trkulja et al. 2013). The virus-infected gloxinia plants exhibit symptoms of chlorotic ringspots and chlorotic and necrotic patterns followed by necrosis and distortion of leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Bertaccini A, Bellardi MG (1990) Tomato spotted wilt virus infecting gloxinia in Italy. Phytopathol Mediterr 29:205–208 Brown LG (1988) Tomato spotted wilt virus in ornamentals. Plant Pathology Circular No.313. Fla. Dept. Agric & Consumer Serv., Division of Plant Industry, Tallahassee, Florida, USA. Chen C-C, Huang CW, Kuo YW, Liu FL, Yuan CH, Hsu HT, Yeh SD (2006) Identification of common epitopes on a conserved region of NSs proteins among tospoviruses of Watermelon silver mottle virus serogroup. Phytopathology 96:1296–1304 Chen C-C, Chang C-A, Yeh S-D, Chen T-C (2012) Occurrence of Capsicum chlorosis virus on ornamental crops in Taiwan. Bioforsk Fokus 7(9):42 Daughtrey ML, Jones RK, Moyer JW, Daub ME, Baker JR (1997) Tospoviruses strike the greenhouse industry: INSV has become a major pathogen on flower crops. Plant Dis 81(11):1220–1230 Hsu H-T, Ueng PP, Chu F-H, Ye Z, Yeh S-D (2000) Serological and molecular characterization of a high temperaturerecovered virus belonging to Tospovirus serogroup IV. J Gen Plant Pathol 66:167–175 Jones RK, Moyer JW (1986) Tomato spotted wilt virus in gloxinia in North Carolina. NC. Flower Growers Bull 30:11–13 Nielsen SL, Nicolaisen M (2010) First report of Columnea latent viroid (CLVd) in Gloxinia gymnostoma, G. nematanthodes and G. purpurascens in a botanical garden in Denmark. New Dis Rep 22:4 Trkulja V, Salapura JM, Curkovic B, Bulajic SA, Vucurovic A, Krstic B (2013) First report of Tomato spotted wilt virus on Gloxinia in Bosnia and Herzegovina. Plant Dis 97(3):429 Zettler FW, Nagel J (1983) Infection of cultivated Gesneriads by two strains of Tobacco mosaic virus. Plant Dis 67:1123–1125

Glycine clandestina (Twining glycine) Family: Fabaceae

Glycine mosaic virus Taxonomic position Genus: Comovirus

Weed host

(GMV)

Family: Secoviridae

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Glycine latifolia (Peak Downs clover)

Geographical distribution GMV infection was first reported in plants of Glycine clandestina from the Blue Mountains, NSW, Australia, by Bowyer et al. (1980). The virus spreads in Australia. Symptoms and host(s) The virus-infected twining glycine plants exhibit symptoms of green mosaic, mottling, and leaf malformation. Transmission The virus is transmitted by beetle vectors. The virus is also transmissible by mechanical sapinoculation. Virion properties and genome The virions are isometric, non-enveloped of two types, but similar in size 28 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is bipartite, of linear positive-sense, singlestranded RNA (Sanfacon et al. 2009; Bruening and Lomonossoff 2011; Sanfacon 2015; Thompson et al. 2017).

References Bowyer JW, Dale JL, Behncken GM (1980) Glycine mosaic virus: a comovirus from Australian native Glycine species. Ann Appl Biol 95:385–390 Bruening G, Lomonossoff GP (2011) Comovirus. Comoviridae. In: The Springer index of viruses. Springer, New York, pp 345–353. https://doi.org/10.1007/978-0-387-95919-1_53 Sanfacon H (2015) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. In: eLS. Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000764.pub3. http://www.els.net Sanfacon H, Wellink J, Le Gall O, Karasev A, van der Vlugt R, Wetzel T (2009) Secoviridae: a proposed family of plant viruses within the order Picornavirales that combines the families Sequiviridae and Comoviridae, the unassigned genera Cheravirus and Sadwavirus, and the proposed genus Torradovirus. Arch Virol 154(5):899–907 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Secoviridae. J Gen Virol 98:529–531

Glycine latifolia (Peak Downs clover) Family: Fabaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Fodder crop

(AMV)

Family: Bromoviridae

AMV infection in plants of Glycine latifolia was reported from Queensland (Australia) (Horlock et al. 1997). The virus-infected Peak Downs clover plants exhibit yellow mosaic symptoms. The virus is transmitted by several species of aphids in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

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References Horlock CM, Teakle DS, Jones RM (1997) Natural infection of the native pasture legume, Glycine latifolia, by Alfalfa mosaic virus in Queensland. Australas Plant Pathol 26:115–116

Glycine max (Soybean) Family: Fabaceae

Oilseed

African cassava mosaic virus Taxonomic position Genus: Begomovirus

(ACMV)

Family: Geminiviridae

ACMV infection in plants of Glycine max was reported from Nigeria and South Africa (Pietersen and Garnett 1990; Alabi et al. 2008; Mgbechi-Ezeri et al. 2008). The virus-infected soybean plants exhibit yellow mosaic and mottling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ACMV, refer to Manihot esculenta.

Ageratum yellow vein virus

(AYVV)

Synonyms Soybean crinkle leaf virus (SbCrLV) Taxonomic position Genus: Begomovirus

Family: Geminiviridae

SbCrLV infection in plants of Glycine max was reported from Thailand (Iwaki et al. 1983; Samretwanich et al. 2001). The virus-infected soybean leaves are twisted or curled, and the undersides have veinal enations. The foliage of diseased plants is darker green than that of healthy plants. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner (Iwaki et al. 1983). The virus is not transmissible by mechanical inoculation; by grafting; nor by seed. For more details of AYVV, refer to Ageratum spp.

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

(AMV)

Family: Bromoviridae

AMV infection in plants of Glycine max was distributed worldwide, including Argentina, Iran, Brazil, Japan, Romania, South Africa, and the USA (Golnaraghi et al. 2002; Giesler and Ziems 2006; Mueller and Grau 2007; Naghavi et al. 2008; Fajolu et al. 2010; He et al. 2011; Hobbs et al. 2012). Infection of

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Glycine max (Soybean)

soybean plants with AMV early in the season result in the mottling or chlorotic spotting of diseased leaves. Later infections result in a yellow mottle, star-shaped flecks or slight crinkling. Out of the 13 aphid vectors transmitting this disease, Myzus persicae is the most efficient vector, and the virus-vector relationship is of the non-persistent type. The virus is mechanically sap-transmissible to certain solanaceous plants and some members of the Leguminosae. The virus is seed-transmitted through soybean seeds up to 9% (He et al. 2010). The virus is not transmissible by contact between plants but is transmissible by grafting. For more details of AMV, refer to Medicago sativa.

Bean common mosaic virus

(BCMV)

Synonyms Blackeye cowpea mosaic virus (BlCMV); Peanut stripe virus (PStV) Taxonomic position Genus: Potyvirus

Family: Potyviridae

BCMV infection in plants of Glycine max was reported from South Korea, China, India, the USA, Taiwan, Korea, and Iran (Dijkstra et al. 1987; Green and Lee 1989; Gillaspie and Hopkins 1991; Vetten et al. 1992; Ghosh and Dhingra 1993; Hristova et al. 1995; Golnaraghi et al. 2002, 2004; Naghavi et al. 2008; Lim 2013; Zhou et al. 2014; Lee et al. 2015; Latha et al. 2017). The virus-infected soybean plants exhibit mosaic, mottling, chlorotic, dark green discoloration, and yellowing symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of BCMV, refer to Phaseolus vulgaris.

Bean golden mosaic virus Taxonomic position Genus: Begomovirus

(BGMV)

Family: Geminiviridae

BGMV infection in plants of Glycine max was reported from Brazil (Fernandes et al. 2009; InoueNagata et al. 2016). The virus-infected soybean plants exhibit bright yellow mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by contact between plants. For more details of BGMV, refer to Phaseolus vulgaris.

Bean pod mottle virus Taxonomic position Genus: Comovirus

(BPMV)

Family: Secoviridae

BPMV infection in plants of Glycine max was first reported from Arkansas (Walters 1958). The virus spreads in Iran, Ecuador, Canada, Alabama, South America, South Africa, and the USA (Lin and Hill 1983; Ross and Butler 1985; Zettler et al. 1989; Ghabrial et al. 1990; Pietersen and Garnett 1990; Anjos et al. 1999; Dorrance et al. 2001; Giesler et al. 2002; Gu et al. 2002; Michelutti et al. 2002; Shahraeen et al. 2005; Sikora and Murphy 2005; Giesler and Ziems 2006). The virus-infected soybean plants

Glycine max (Soybean)

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exhibit symptoms of crinkling, mottling, leaf distortion, blistering of leaves, and stunting of plants; some infected soybean plants had green stems and undersized and “turned up pods.” The virus is transmitted by the bean leaf beetle vector, Cerotoma trifurcata, in a non-persistent manner (Bradshaw et al. 2007). This virus is transmitted through seed of soybean at very low rate of 0.1% (Lin and Hill 1983; Mabry et al. 2003; Yi et al. 2011). For more details of BPMV, refer to Phaseolus vulgaris.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Glycine max spreads in the USA, Argentina, Brazil, the former USSR, Iran, and Asia (Pietersen and Garnett 1990; Golnaraghi et al. 2000, 2002, 2004; Campos et al. 2014; Sharma et al. 2016). The virus-affected younger leaves on soybean plants show a scattered yellow mottling or indefinite yellow bands along the major veins. Usually, the leaves are not distorted and puckered like those affected by soybean mosaic, but some strains of BYMV produce severe mottling and crinkling of leaves. Affected plants are not noticeably stunted. Rusty-appearing dead spots develop in the yellowed areas as the leaves mature. Some strains produce severe mottling and crinkling of leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Blackgram mottle virus

(BMoV)

Taxonomic position BMoV is a tentative member of the genus Carmovirus. BMoV infection in plants of Glycine max was reported from India and Thailand (Honda et al. 1982; Sharma et al. 2016). The virus-infected soybean plants exhibit mottling and mosaic symptoms. The virus is transmitted by beetle vectors and is also transmissible by mechanical sap-inoculation. For more details of BMoV, refer to Vigna mungo.

Brome mosaic virus Taxonomic position Genus: Bromovirus

(BMV)

Family: Bromoviridae

BMV infection in plants of Glycine max was reported from Canada (Diaz-Cruz et al. 2018). The virusinfected soybean plants exhibit symptoms of dark spots with chlorotic halos. The virus is transmitted by the cereal leaf beetle vector, Oulema melanopus, and is also transmissible by mechanical sap-inoculation to many monocotyledonous and a few dicotyledonous plants. For more details of BMV, refer to Bromus spp.

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Glycine max (Soybean)

Chino del tomate virus Taxonomic position Genus: Begomovirus

(CdTV)

Family: Geminiviridae

CdTV infection in plants of Glycine max was reported from Mexico (Mauricio-Castillo et al. 2014). The virus-infected soybean plants exhibit dwarfing, rolling, and yellowing of leaves. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by mechanical inoculation nor by contact between plants. For more details of CdTV, refer to Solanum lycopersicum.

Clover yellow vein virus Taxonomic position Genus: Potyvirus

(ClYVV)

Family: Potyviridae

ClYVV infection in plants of Glycine max was reported from Korea (Shin et al. 2014). The virusinfected soybean plants exhibit mosaic and mottling symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ClYVV, refer to Trifolium spp.

Cotton leaf curl Kokhran virus Taxonomic position Genus: Begomovirus

(CLCuKoV)

Family: Geminiviridae

CLCuKoV infection in plants of Glycine max was reported from India (Raj et al. 2006a). The virusinfected soybean plants exhibit yellow mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is neither transmissible by mechanical sap-inoculation, and also by contact between plants. For more details of CLCuKoV, refer to Gossypium spp.

Cowpea aphid-borne mosaic virus Taxonomic position Genus: Potyvirus

(CABMV)

Family: Potyviridae

CABMV infection in plants of Glycine max was reported from Ceara (Brazil) (Sousa et al. 1996). The virus-infected soybean plants exhibit systemic mosaic symptoms. The virus is transmitted by the aphid vector, Aphis gossypii, in a non-persistent manner, and is also transmissible by mechanical sapinoculation. The virus is transmitted by cowpea seeds to the extent of 4.1% (Sousa et al. 1996). For more details of CABMV, refer to Vigna unguiculata.

Glycine max (Soybean)

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Cowpea chlorotic mottle virus Taxonomic position Genus: Bromovirus

(CCMV)

Family: Bromoviridae

CCMV infection in plants of Glycine max was reported from the USA and South Africa (Kuhn 1968; Pietersen and Garnett 1990). The virus-infected soybean plants leaves are mottled, slightly crinkled, and tend to stand upright. Mottling is the most intense on the diseased younger leaves. Plants are stunted and produce fewer seeds, which are small in size and reduced in quality. The virus is transmitted by the bean leaf beetle (Cerotoma trifurcata) and the spotted cucumber beetle (Diabrotica undecimpunctata) and is also transmissible by mechanical sap-inoculation. For more details of CCMV, refer to Vigna unguiculata.

G Cowpea mild mottle virus Taxonomic position Genus: Carlavirus

(CPMMV)

Family: Betaflexiviridae

CPMMV infection in plants of Glycine max was distributed in Asia, Ivory Coast, Thailand, Egypt, Argentina, Brazil, and certain African countries (Iwaki et al. 1982, 1986a; Thouvenel et al. 1982; Anno Nyako 1984; El-Hammady et al. 2004; Almeida et al. 2005a; Laguna et al. 2006; Tavassoli et al. 2008, 2009; Yadav et al. 2013a, b; Zanardo et al. 2014; Zinsou et al. 2015; Sharma et al. 2016; Zanardo and Carvalho 2017). The virus-infected soybean plants are stunted and show light green and yellow mosaic. Leaves may also be curled downward or cupped upward. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a semi-persistent manner (Iwaki et al. 1982; Marubayashi et al. 2010) and is also transmissible by mechanical sap-inoculation. The virus is seed-transmitted from 0.5% to 0.9% through infected soybean seeds (Iwaki et al. 1982; Thouvenel et al. 1982). For more details of CPMMV, refer to Vigna unguiculata.

Cowpea mosaic virus Taxonomic position Genus: Comovirus

(CPMV)

Family: Secoviridae

CPMV infection in plants of Glycine max was reported from Central America, the USA, and Korea (Thongmeearkom and Goodman 1976; McLaughlin et al. 1978; Cho and Lee 2003). Inoculated primary leaves have chlorotic and necrotic lesions. A systemic light green mosaic occurs on diseased trifoliate leaves together with very severe leaf malformation and bud blight. The virus is transmitted by the bean leaf beetle vector, Cerotoma ruficornis, and is also transmissible by mechanical sap-inoculation. For more details of CPMV, refer to Vigna unguiculata.

Cowpea mottle virus Taxonomic position Genus: Gammacarmovirus

(CPMoV)

Family: Tombusviridae

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Glycine max (Soybean)

CPMoV infection in plants of Glycine max was reported from Nigeria (Arogundade et al. 2010). The virus-infected soybean plants exhibit leaf mottling, which progresses to leaf wrinkling and affects fruit setting. The virus is transmitted by beetle vectors and is also transmissible by mechanical sapinoculation. For more details of CPMoV, refer to Vigna unguiculata.

Cowpea severe mosaic virus Taxonomic position Genus: Comovirus

(CPSMV)

Family: Secoviridae

CPSMV infection in plants of Glycine max was reported from Illinois, USA and Brazil (Anjos and Lin 1984). The virus-infected soybean plants exhibit typical bud blight symptoms. The virus causes severe mosaic and stunting. The virus is transmitted by numerous leaf-feeding beetle vectors such as Cerotoma arcuata, and is also transmissible by mechanical sap-inoculation. For more details of CPSMV, refer to Vigna unguiculata.

Cucumber mosaic virus

(CMV)

Synonyms Soybean stunt virus

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV infection in plants of Glycine max was reported from India, Iran, Nigeria, and Japan (Hanada and Tochihara 1982; Golnaraghi et al. 2002, 2004; Hong et al. 2003, 2007; Naghavi et al. 2008; Arogundade et al. 2010; Hosseinzadeh et al. 2012; Sharma et al. 2016). The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Groundnut bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(GBNV)

Family: Tospoviridae

GBNV infection in plants of Glycine max occurs worldwide (Bhat et al. 2002a; Kumari et al. 2003; Sharma et al. 2016). The characteristic symptoms on soybean include necrosis of plant parts such as leaves, stems, petioles, growing points, buds, and pods. Plants infected early in the season are severely stunted with reduced intermodal length and many axillary shoots. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a number of hosts. For more details of GBNV, refer to Arachis hypogaea.

Glycine max (Soybean)

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Groundnut ringspot orthotospovirus Taxonomic position Genus: Orthotospovirus

(GRSV)

Family: Tospoviridae

GRSV infection in plants of Glycine max was reported from South Africa (Pietersen and Staples 1996; Pietersen and Morris 2002). The virus-infected soybean plants exhibit leaf mottle symptoms. The virus is transmitted by the thrips vectors, F. occidentalis and F. schultzei, in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of GRSV, refer to Arachis hypogaea.

Indonesian soybean dwarf virus Taxonomic position Genus: Unassigned

(ISDV)

Family: Luteoviridae

Geographical distribution ISDV infection in plants of Glycine max was reported from Indonesia and Thailand (Iwaki et al. 1980). Symptoms and host(s) The virus-infected soybean plants are dwarfed and dark green with shortened leaf petioles and internodes. Upper leaves are small and curl upward. Lower leaves are rugose and brittle and have shortened veins and interveinal white necrosis. Occasionally diseased leaves have numerous holes caused by their brittleness. Diseased plants produce few pods. Transmission The virus is transmitted by the aphid vector, Aphis glycines, in a persistent manner. The virus is retained when the vector molts, does not multiply in the vector, and is not transmissible by mechanical inoculation, by contact between plants, by seed, and by pollen. Virion properties and genome The virions are isometric, non-enveloped, 26 nm in diameter, rounded in profile, and without a conspicuous capsomere arrangement.

Macroptilium yellow spot virus

(MacYSV)

Synonyms Soybean chlorotic spot virus (SoCSV) Taxonomic position Genus: Begomovirus

Family: Geminiviridae

SoCSV infection in plants of Glycine max was reported from Brazil (Coco et al. 2013). The virusinfected soybean plants exhibit mild symptoms of chlorotic spots on the leaves. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of MacYSV, refer to Macroptilium spp.

G

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Mungbean yellow mosaic India virus Taxonomic position Genus: Begomovirus

(MYMIV)

Family: Geminiviridae

MYMIV infection in plants of Glycine max was reported from India, Indonesia (Usharani et al. 2004, 2005; Girish and Usha 2005; Tsai et al. 2013; Sharma et al. 2016). The virus-infected soybean plants exhibit yellow mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by mechanical inoculation. For more details of MYMIV, refer to Vigna radiata.

Mungbean yellow mosaic virus Taxonomic position Genus: Begomovirus

(MYMV)

Family: Geminiviridae

MYMV infection in plants of Glycine max is present in Asian countries including India and Thailand (Usharani et al. 2004; Girish and Usha 2005). The virus-affected soybean plants show faint yellow specks or spots and irregular blotching of tip leaves, with slight leaf malformation. Some cultivars produce typical yellow and golden mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by mechanical inoculation. For more details of MYMV, refer to Vigna radiata.

Okra mottle virus Taxonomic position Genus: Begomovirus

(OMoV)

Family: Geminiviridae

OMoV infection in plants of Glycine max was reported from Brazil (Fernandes et al. 2009). The virusinfected soybean plants exhibit leaf mottling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of OMoV, refer to Abelmoschus esculentus.

Papaya leaf crumple virus Taxonomic position Genus: Begomovirus

(PaLCrV)

Family: Geminiviridae

PaLCrV infection in plants of Glycine max was reported from India (Jaidi et al. 2015). The virusinfected soybean plants exhibit yellow mosaic on leaves and leaf crumpling and distortion symptoms. Severely infected plants were stunted and bore fewer number of flowers and pods. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of PaLCrV, refer to Carica papaya.

Glycine max (Soybean)

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Pea enation mosaic virus 1 Taxonomic position Genus: Enamovirus

(PEMV-1)

Family: Luteoviridae

PEMV-1 infection in plants of Glycine max was reported from Iran (Golnaraghi et al. 2002, 2004). The virus-infected soybean plants exhibit mosaic or mottling symptoms. The virus is transmitted by aphid vectors in a circulative, non-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of PEMV-1, refer to Pisum sativum.

Peanut mottle virus Taxonomic position Genus: Potyvirus

(PeMoV)

G Family: Potyviridae

PeMoV infection in plants of Glycine max was reported from southeastern USA, Africa, northeast Australia, West Kenya, Uganda, Venezuela, Malaysia, Bulgaria, South Korea, Iran, and Asian countries (Kuhn et al. 1972; Demski 1975; Demski and Kuhn 1977; Roane et al. 1978; Pietersen and Garnett 1992; Golnaraghi et al. 2000, 2002, 2004; Lim et al. 2014). The virus-infected soybean plants exhibit systemic chlorotic mottle, mild mosaic or mild to severe leaf curling, and distortion. The virus is transmitted by several aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. Seed transmission was up to 0.22% through PeMoV-infected soybean seeds (Iwaki et al. 1986b). For more details of PeMoV, refer to Arachis hypogaea.

Peanut stunt virus Taxonomic position Genus: Cucumovirus

(PSV)

Family: Bromoviridae

PSV infection in plants of Glycine max was reported from the southeastern and central USA and Japan (Iizuka and Yunoki 1974; Milbrath and Tolin 1977). The virus-infected soybean plants exhibit symptoms ranging from diffuse chlorotic lesions to necrotic local lesions on inoculated primary leaves. Vein-clearing occurs, followed by a general mosaic of diseased trifoliate leaves. Some diseased plants may be symptomless. The virus is transmitted by aphids in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. The virus is seed-transmitted through an infected soybean seed of up to 0.2% (Iizuka 1973). For more details of PSV, refer to Arachis hypogaea.

Pepper leafroll virus Taxonomic position Genus: Begomovirus

(PepLRV)

Family: Geminiviridae

PepLRV infection in plants of Glycine max was reported from Ecuador (Fiallo-Olivé et al. 2019). The virus-infected soybean plants show leaf deformation and/or mosaic symptoms. The virus is transmitted

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Glycine max (Soybean)

by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner. For more details of PepLRV, refer to Capsicum baccatum cv. Pendulum.

Quail pea mosaic virus Taxonomic position Genus: Comoviru

(QPMV)

Family: Secoviridae

QPMV infection in plants of Glycine max was reported from Venezuela (Debrot and Centeno 1986). The virus-infected soybean plants exhibit mosaic symptoms. The virus is transmitted by beetle vectors and is also transmissible by mechanical sap-inoculation. For more details of QPMV, refer to Strophostyles helvola.

Rhynchosia golden mosaic virus Taxonomic position Genus: Begomovirus

(RhGMV)

Family: Geminiviridae

RhGMV infection in plants of Glycine max was reported from Mexico (Mendez-Lozano et al. 2006). The virus-infected soybean plants exhibit symptoms of yellowing, curled leaves, and stunting. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of RhGMV, refer to Rhynchosia minima.

Sida micrantha mosaic virus Taxonomic position Genus: Begomovirus

(SiMMV)

Family: Geminiviridae

SiMMV infection in plants of Glycine max was reported from Brazil (Fernandes et al. 2009). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of SiMMV, refer to Sida spp.

Southern bean mosaic virus Taxonomic position Genus: Sobemovirus

(SBMV)

Family: Solemoviridae

SBMV infection in plants of Glycine max was reported from China (Iizuka and Yunoki 1975; Xu et al. 1986; Pietersen and Garnett 1990). The virus-infected soybean plants show mild systemic mottle with most strains depending on variety. The virus is transmitted by beetle vectors in a semi-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of SBMV, refer to Phaseolus vulgaris.

Glycine max (Soybean)

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Soybean blistering mosaic virus Taxonomic position Genus: Begomovirus

(SbBMV)

Family: Geminiviridae

Geographic distribution SbBMV infection in plants of Glycine max was reported from Argentina (Rodriguez-Pardina et al. 2011). Symptoms and host(s) The virus-infected soybean plants exhibit a foliar yellow mosaic and blistering symptoms on the leaves. Transmission Insect transmission of SbBMV has not been demonstrated. However, in common with all other begomoviruses, SbBMV is likely transmitted by the whitefly vector of begomoviruses, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of SbBMV has not been investigated. In common with all geminiviruses, the virions of SbBMV are likely geminate (twinned icosahedra). It is unclear at this time whether SbBMV is a bipartite begomovirus, as is typical of the majority of begomoviruses native to the New World, or monopartite since only the sequence of a single-genome/ DNA-A genomic component has been submitted to the nucleotide sequence databases (EF016486 = NC_038463). Only one monopartite begomovirus has been confirmed to occur in the New World: Tomato leaf deformation virus (Melgarejo et al. 2013; Sánchez-Campos et al. 2013). The available sequence of SbBMV is a circular molecule of 2605 nt (EF016486) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). Typical of the DNA-A component of bipartite begomoviruses and the genomes of monopartite begomoviruses native to the New World, the sequence is predicted to encode five genes, one in the virion-sense and four in the complementary-sense. The expression and function of these genes have not been investigated for SbBMV.

Soybean blotchy mosaic virus (SbBMV) Taxonomic position SbBMV is a tentative member of the genus Cytorhabdovirus and family Rhabdoviridae Geographical distribution SbBMV-infected Glycine max plants were reported from South Africa by Lamprecht et al. (2010). Symptoms and host(s) The virus-infected soybean plants displayed distinct blotchy mosaic symptoms on the leaves. Transmission The virus is transmitted by the leafhopper vector Peragallia caboverdensis in a persistent, propagative manner. Mechanically transmissible from soybean to soybean with transmission frequency less than 30%. Mechanical transmission from soybean to N. benthamiana and between individual N. benthamiana plants resulted in symptomless-infected plants but at transmission rates of 30–40%.

G

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Glycine max (Soybean)

Virion properties and Genome The virions are bullet-shaped morphology and 338–371 nm in length and 93 nm in wide. A partial genome sequence of 522 nt is available (EU877231) (Walker et al. 2018).

Soybean chlorotic blotch virus Taxonomic position Genus: Begomovirus

(SbCBV)

Family: Geminiviridae

Geographical distribution SbCBV infection in plants of Glycine max was reported from Nigeria (Alabi et al. 2010). Symptoms and host(s) The virus-infected soybean plants exhibit chlorotic blotches and stunting symptoms. The virus has been isolated from soybean (Glycine max), Centrosema pubescens, and lima bean (Phaseolus lunatus). Transmission Insect transmission of SbCBV has not been demonstrated. However, in common with all other begomoviruses, SbCBV is likely transmitted by the whitefly vector of begomoviruses, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of SbCBV has not been investigated. In common with all geminiviruses, the virions of SbCBV are likely geminate (twinned icosahedra). SbCBV is a typical bipartite begomovirus native to the Old World. The genome consists of two circular, single-stranded DNA components. DNA-A contains 2708 nt (GQ472985 = NC_014141; GQ472987) and DNA-B of 2647 nt (GQ472986 = NC_014142; GQ472988) (Briddon 2001; Alabi et al. 2010; Brown et al. 2015; Zerbini et al. 2017). Typical of the majority of bipartite begomoviruses native to the Old World, the DNA-A component of SbCBV encodes six genes, two in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for SbCBV.

Soybean chlorotic mottle virus Taxonomic position Genus: Soymovirus

(SbCMV)

Family: Caulimoviridae

Geographical distribution SbCMV infection in plants of Glycine max was reported from Japan (Iwaki et al. 1984; Hibi and Kameya-Iwaki 1988). Symptoms and host(s) The virus-infected soybean plants exhibit vein-clearing and chlorosis on young leaves which develop into mottle and mosaic symptoms. Plants are stunted due to shortened internodes.

Glycine max (Soybean)

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Transmission The virus is transmissible by mechanical sap-inoculation. Inoculated Dolichos lablab, Phaseolus vulgaris, and Vigna unguiculata show various degrees of systemic symptoms. The virus is not transmitted by aphid vectors, nor by seed. Virion properties and Genome The virions are isometric, non-enveloped, and 50 nm in diameter with no obvious surface structure. The genome is a monopartite, circular, double-stranded DNA of 8178 bp (X15828 = NC_001739) (Hibi et al. 1986; Verver et al. 1987; Hasegawa et al. 1989; Takemoto and Hibi 2001; Hibi 2011).

Soybean dwarf virus Taxonomic position Genus: Luteovirus

(SbDV)

G Family: Luteoviridae

Geographical distribution SbDV was first reported in plants of Glycine max in Japan by Tamada et al. (1969). In due course, this virus was reported in Indonesia, Africa, Australia, Japan, New Zealand, and in California (USA) (Takahashi et al. 1980; Rossel and Thottappilly 1983; Damsteegt et al. 1990; Phibbs et al. 2004; Harrison et al. 2005; Thekkeveetil et al. 2007; Hobbs et al. 2012). Symptoms and host(s) The virus-infected soybean plants are severely stunted. The leaves are rugose, curl downward, and have interveinal yellowing. Terauchi et al. (2001) have reported four strains of SbDV (YS, YP, DS, and DP) based on symptomatology on infected soybeans and aphid vector specificity. YS and YP strains cause yellowing in soybean and infect white clover (Trifolium repens), but not red clover (T. pratense). DS and DP strains cause dwarfing in soybeans and infect red clover but not white clover (Damsteegt et al. 1990). The natural host range of this virus includes Pisum sativum, Trifolium spp., Phaseolus vulgaris, Lupinus spp., Medicago spp., and V. faba (Tamada 1970). Transmission The virus is transmitted by the aphid vector Acyrthosiphon solani (Aulacorthum) (and some isolates also by Acyrthosiphon pisum) in a circulative, non-propagative manner (Damsteegt and Hewings 1986; Harrison et al. 2005). Virus is retained when the vector molts, does not multiply in the vector, is not transmitted congenitally to the progeny of the vector, and does not require a helper virus for vector transmission. YS and DS strains are transmitted by the aphid vector Aulacorthum solani, while YP and DP strains are transmitted by both Acyrthosiphon pisum and Nearctaphis bakeri (Honda et al. 1999). The virus is not mechanically sap-transmissible. The virus is not transmissible by contact between plants and not transmitted by seed (Tamada 1975). Virion properties and genome The virions are 25–30 nm in diameter, hexagonal in outline, and are non-enveloped (Tamada 1975; Kojima and Tamada 1976). The genome is a monopartite, linear, positive-sense, single-stranded RNA of 5853 nt (NC_003056). The 30 terminus has neither a poly (A) tract nor a tRNA-like structure. There is no genome-linked protein (VPg) at the 50 -terminus. The genome consists of five or six ORFs that are

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Glycine max (Soybean)

predicted to encode proteins of between 4 and 132 kDa (Hewings et al. 1986; Rathjen et al. 1994; Terauchi et al. 2001).

Soybean latent spherical virus

(SLSV)

Taxonomic position Genus: Nepovirus Family: Secoviridae Geographical distribution SLSV infection in plants of Glycine max was reported from the USA (Yasmin et al. 2017). Transmission The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sapinoculation, and by grafting. Virion properties and genome Virions are isometric, non-enveloped of two types but similar in size 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is bipartite, linear positive-sense, singlestranded RNA. RNA1 contains 8170 nt (KX424571) and RNA2 of 5776 nt (KX424572 = NC_032271) (Sanfacon 2015; Thompson et al. 2017; Yasmin et al. 2017).

Soybean leaf rugose mosaic virus

(SLRMV)

Taxonomic position SLRMV is a tentative member of the genus Bymovirus and family Potyviridae Geographical distribution SLRMV infection in plants of Glycine max was reported from Japan (Kuroda et al. 2010). Symptoms and host(s) The virus-infected soybean plants exhibit stunting, severe mosaic, and leaf rugose symptoms. Transmission The virus is mechanically sap-transmissible to Gomphrena globosa causing necrotic spots. The virus is not transmitted by the aphid vectors, Aphis craccivora, A. glycines, or Aulacorthum solani. The virus is not seed-transmitted (Kuroda et al. 2010). When soybeans are grow in virus-infested soil, the plants became virus-infected indicating the involvement of soil-borne vector. Virion properties and genome The virions are flexuous and filaments virus-like particles with a modal length of 500 nm. The genome is single-stranded RNA. A partial genome sequence of 1435 nt is available (AB560671) (Kuroda et al. 2010; Wylie et al. 2017).

Glycine max (Soybean)

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Soybean mild mottle virus Taxonomic position Genus: Begomovirus

(SbMMoV)

Family: Geminiviridae

Geographical distribution SbMMoV infection in plants of Glycine max was reported from Nigeria (Alabi et al. 2010). Symptoms and host(s) The virus-infected soybean plants exhibit mild foliar yellow mosaic symptoms. Transmission Insect transmission of SbMMV has not been demonstrated. However, in common with all other begomoviruses, SbMMV is likely transmitted by the whitefly vector of begomoviruses, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of SbMMV has not been investigated. In common with all geminiviruses, the virions of SbMMV are likely geminate (twinned icosahedra). The genome of SbMMV consists of a single, circular, single-stranded DNA of 2768 nt (GQ472984 = NC_014140) (Briddon 2001; Alabi et al. 2010; Brown et al. 2015; Zerbini et al. 2017). The characterized genome of SbMMV encodes the six genes, typically encoded by monopartite begomoviruses and the DNA-A component of bipartite begomoviruses. The expression and function of the genes have not been investigated for SbMMV.

Soybean mosaic virus Taxonomic position Genus: Potyvirus

(SMV)

Family: Potyviridae

Geographical distribution SMV was first documented in the USA in 1915 by Clinton (1916), and SMV was named by Gardner and Kendrick (1921). The virus infection of Glycine max plants is worldwide in distribution including Japan, Southeast Asia, China, Iran, Argentina, Poland, Korea, Brazil, South Korea, and the USA (Takahashi et al. 1980; Chen et al. 1986; Xu et al. 1986; Hill et al. 1987; Fiedorow 1994; Almeida et al. 1995; Zhou et al. 1995; Zheng et al. 2000; Saruta et al. 2005; Giesler and Ziems 2006; Naghavi et al. 2008; Nelson and Domier 2009; Li et al. 2010; Lu et al. 2010; Cho et al. 2013; Banerjee et al. 2014; Seo et al. 2014; Hema et al. 2015; Jezewska et al. 2015; Li et al. 2015). Symptoms and host(s) The virus-infected soybean plants exhibit symptoms of characteristic mosaic pattern of alternate light and dark green patches on the leaves. The leaves become leathery rugose and become brittle as diseased plants mature. Plants infected early in the season are stunted, have shortened petioles and internodes, and often show a browning of stems and petioles. Typically the infected plants mature conspicuously later than uninfected ones, in the field the remain green, while most of the other plants become defoliated and dried (Liao et al. 1995). Numerous SMV isolates have been reported worldwide. The isolates from Japan were classified into five strains A, B, C, D, and E based on symptoms induced on

G

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Glycine max (Soybean)

four soybean cultivars. Similarly in China eight strains were isolated (Cho and Goodman 1979; Anjos et al. 1985; Jain et al. 1992; Guo et al. 2005; Viel et al. 2009). Transmission The virus is aphid transmitted, and nearly 31 species are involved, viz., Acyrthosiphon pisum, Aphis craccivora, A. fabae, A. glycine, A. gossypii, Myzus persicae, Rhopalosiphum maidis, and R. padi, in a non-persistent manner (Abney et al. 1976; Lucas and Hill 1980; Hill et al. 2001; Clark and Perry 2002; Domier et al. 2003; Damsteegt et al. 2011). The virus is mechanically sap-transmissible and has a relatively narrow host range. It infects species in six plant families, i.e., Fabaceae (also Leguminosae), Amaranthaceae, Chenopodiaceae, Passifloraceae, Scrophulariaceae, and Solanaceae, but mostly the Leguminosae including soybean and its wild relatives (Hill 1999). The virus is seed-transmitted in soybean at rates in the range of 7–52% (Goodman and Oard 1980; Bowers and Goodman 1979, 1991; Almeida et al. 1995; Pacumbaba 1995; Naik and Murthy 1997; Chalam et al. 2004; Bashar 2015). Virion properties and genome The virions are flexuous filaments and non-enveloped, with a clear modal length of 650–760 nm and 15–18 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of about 9588 nt encoding ten different functional and structural proteins (D00507 = NC_002634). The molecular weights of the RNA and coat protein (the only structure protein) are 3.25  106 and 28,300 Da, respectively (Galvez 1963; Jayaram et al. 1992; El Ahdal 1996; Hill 1999; Chen et al. 2002; Lim et al. 2003; Seo et al. 2009; Zhang et al. 2012; Revers and Garcia 2015; Wylie et al. 2017).

Soybean Putnam virus Taxonomic position Genus: Caulimovirus

(SPuV)

Family: Caulimoviridae

Geographical distribution SPuV infection in plants of Glycine max was reported from Putnam County, Ohio, USA (Han et al. 2012). Transmission The virus is transmitted by aphid vectors in a semi-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are isometric, non-enveloped, and 45–50 nm in diameter with no obvious surface structure. The genome is a monopartite, circular, double-stranded DNA of 7757 bp (JQ926983) (Hohn 2011; Han et al. 2012).

Soybean vein necrosis associated virus

(SVNaV)

Taxonomic position SVNaV is a tentative member of the genus Orthotospovirus and family Tospoviridae Geographical distribution SVNaV infection in plants of Glycine max was reported from Ohio, Oklahoma, Wisconsin, Iowa, and Alabama (USA), and Canada (Zhou 2012; Conner et al. 2013; Han et al. 2013; Smith et al. 2013).

Glycine max (Soybean)

1091

Symptoms and host(s) The virus-infected soybean plants exhibit symptoms including systemic mosaic accompanied by leaf deformation, chlorosis, vein necrosis, and rusty spots on mature leaves. In severe cases the leaves will dry off. Transmission The virus is transmitted by a thrips vector, Sericothrips variabilis, in a persistent-propagative manner. The virus is not seed-transmitted (Hajimorad et al. 2015). Virion properties and genome The genome consists of a 16.5 kb tripartite ambisense, single-stranded RNA (Zhou et al. 2011; Khatabi et al. 2012; Zhou and Tzanetakis 2013).

G Soybean vein necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(SVNV)

Family: Tospoviridae

Geographical distribution SVNV infection in plants of Glycine max was reported from at least ten states of the USA, Egypt, and Canada (Tzanetakis et al. 2009; Zhou et al. 2011; Tenuta 2012; Ali and Abdalla 2013; Ali and Mohammad 2013; Jacobs and Chilvers 2013; Bloomingdale et al. 2014; Abd El-Wahab and El-Shazly 2017; Sikora et al. 2017; Chitturi et al. 2018; Zhou and Tzanetakis 2018, 2019). Symptoms and host(s) The virus-infected soybean plants exhibit symptoms of leaf chlorosis and leaf vein necrosis on some soybean plants in the field (Ali and Abdalla 2013). Initially, small light green to yellow patches develop near main leaf veins. These patches then develop into a mottled light green-yellow-brown pattern. The veins in these areas of the leaflet may become clear to almost translucent which is referred to as veinclearing. As the disease progresses, these areas may have a scaly or scabby appearance. On more susceptible varieties, the brown areas may expand killing larger areas of leaf tissue and giving a scorched appearance to the leaves. Transmission The virus is transmitted by several thrips species including Neohydatothrips variabilis (soybean thrips), Frankliniella tritici (eastern flower thrips) and Frankliniella fusca (tobacco thrips) in a persistent, propagative manner (Keough et al. 2016; Chitturi et al. 2018; Han et al. 2019). The virus is mechanically sap-transmissible to Glycine max, Vigna unguiculata, V. radiata, Nicotiana benthamiana, N. tabacum, N. glutinosa, and Cucurbita pepo (Bloomingdale et al. 2014). SVNV is transmitted through soybean (Glycine max) seed to seedlings from an infected seed lot at a rate of 6% (Groves et al. 2016). Virion properties and genome Virions are spherical or pleomorphic, 80 -120 nm in diameter (King et al., 2012). The virus had a genome consisting of three single-stranded RNA molecules, namely the large (L) 9010 nt [HQ728385, GU722317], medium (M) 4955 nt [HQ728387, GU722318, HQ728386] and small (S) 2603 nt [HQ728386, GU722319] RNAs (Zhou et al., 2011).

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Glycine max (Soybean)

Soybean yellow common mosaic virus Taxonomic position Genus: Sobemovirus

(SYCMV)

Family: Solemoviridae

Geographical distribution SYCMV infection in plants of Glycine max was reported from Korea (Nam et al. 2012; Cho et al. 2013; Seo et al. 2014; Li et al. 2015). Symptoms and host(s) The virus-infected soybean plants exhibit yellow mosaic symptoms. Transmission The virus is mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped icosahedral particles with a diameter of 25–33 nm. The genome is a polycistronic positive-sense single-stranded RNA (ssRNA) of 4152 nt (JF495127 = NC_016033). The virus contains four putative open reading frames encoding P1 (78–566 nt), polyprotein ORF2a (524–2248 nt), polymerase domain ORF2b (1852–3417 nt), and CP (3227–4030 nt) (Nam et al. 2012; Somera et al. 2015).

Soybean yellow mottle mosaic virus Taxonomic position Genus: Gammacarmovirus

(SYMMV)

Family: Tombusviridae

Geographical distribution SYMMV infection in plants of Glycine max was reported from Korea, North America, and India (Li et al. 2009; Cho et al. 2013; Seo et al. 2014; Li et al. 2015; Sandra et al. 2015). Symptoms and host(s) The virus-infected soybean plants display bright yellow mosaic symptoms and reduced growth. The natural host range of this virus is soybean, mung bean, black gram, French bean, and guar bean (Lee and Kim 2013; Sandra et al. 2015). Transmission The virus is mechanically sap-transmissible to only Leguminosae members but not to Amaranthaceae, Cucurbitaceae and Solanaceae. Virion properties and genome The virions are isometric 30 nm in diameter and have T = 3 icosahedral symmetry. The genome consists of a single molecule of linear positive-sense ssRNA of 4009 nt (FJ457015 = NC_011643) (Nam et al. 2009).

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Tobacco mosaic virus

(TMV)

Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

TMV-infected plants of Glycine max were reported from Yugoslavia, Iran, Romania, and India (Taraku and Tolin 1986; McDaniel et al. 1995; Golnaraghi et al. 2002; Kumar et al. 2013, 2017). The virusinfected soybean plant leaves exhibit vein-clearing and mild chlorotic mosaic symptoms. Tobacco mosaic virus soybean strain (TMV-S) is mechanically sap-transmissible, and the host range of TMV-S differs from that of TMV. No vector is known for this virus. For more details of TMV, refer to Nicotiana tabacum.

Tobacco necrosis virus A Taxonomic position Genus: Alphanecrovirus

(TNV-A)

Family: Tombusviridae

TNV-A infection in plants of Glycine max was reported from China (Xi et al. 2008). The virus-infected soybean plants exhibit necrotic streaks on the stem and petioles. The virus is transmitted by a fungal vector, Olpidium brassicae, and is also transmissible by mechanical sap-inoculation. For more details of TNV-A, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV occurs in almost all countries that grow Glycine max including Australia, Canada, Cuba, Brazil, Asia, and the USA (Gupta 1978; Almeida 1980; Tu 1986; Xu et al. 1986; Demski and Kuhn 1989; Pietersen and Garnett 1990; Demski et al. 1999; Zhao et al. 2015). The terminal bud of virus-infected young soybean plants turns brown and bends sharply downward forming a crook. Later, other buds become brown. Such buds become dry and brittle and may fall off at the slightest touch. This disease is also known as bud blight. The leaf just below the tip bud may be bronzed and flecked in a rust color. A reddish brown discoloration of the pith, especially at the nodes (joints), is often evident inside the stem. Brown streaks are sometimes evident on petioles and along large leaf veins. Plants infected before flowering time are severely dwarfed and produce few or no pods. The virus is transmitted by the nematode vector, Xiphinema americanum, in a non-persistent manner, and is also transmissible by mechanical sap-inoculation (Brown and Trudgill 1998). The virus is seed-transmitted in soybean to the extent of 90–100% (Iizuka 1973; Demski and Harris 1974; Yi et al. 2011). For more details of TRSV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

G

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Glycine max (Soybean)

TSV infection in plants of Glycine max was reported from Brazil, Argentina, Australia, India, and the USA (Fagbenle and Ford 1970; Sherwood and Jackson 1985; Pietersen and Garnett 1990; Ravi et al. 2001; Bhat et al. 2002b; Rebedeaux et al. 2005; Almeida et al. 2005b; Arun Kumar et al. 2008; Dutta et al. 2015; Sharman et al. 2015; Irizarry et al. 2016; Rajamanickam et al. 2016; Sharma et al. 2016). The characteristic symptoms on soybean produced by TSV are chlorosis, mosaic mottling, necrosis of leaves, petiole stem and pod necrosis, and dieback of terminal portions of stems, commonly called bud blight. Sometimes infected plants partially recover and then develop supernumerary axillary branches which are stunted and produce dwarfed leaves. Necrotic blotches appear on pods (Fagbenle and Ford 1970). Comparison of different TSV isolates from soybean has been studied by Ghanekar and Schwenk (1980). The virus is transmitted by the thrips vectors (Frankliniella occidentalis and Thrips tabaci); the virus present in/on pollen entering the host through injuries caused by thrips while feeding. The virus is transmissible by mechanical sap-inoculation, and plants belonging to more than nine families are susceptible. The virus is seed-transmitted to the extent of 30–90% through soybean seeds (Ghanekar and Schwenk 1974; Truol et al. 1987). The virus is pollen transmitted. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato leaf curl Karnataka virus Taxonomic position Genus: Begomovirus

(ToLCKaV)

Family: Geminiviridae

ToLCKaV infection in plants of Glycine max was reported from India (Raj et al. 2006b). The virusinfected soybean plants exhibit symptoms of severe yellowing, crumpling, and distortion of leaves. The infected plants remained dwarf with fewer flowers which resulted in a very low yield. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner, and also by grafting. For more details of ToLCKaV, refer to Solanum lycopersicum.

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Glycine max was reported from Pakistan (Jamil et al. 2017). The virusinfected soybean plants exhibit severe leaf curling, vein thickening, and leaf yellowing symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is graft-transmissible. For more details of ToLCNDV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Glycine max was reported from Iran (Golnaraghi et al. 2004). The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical

Glycine max (Soybean)

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sap-inoculation, and by grafting. The virus is seed-transmitted in soybean seeds up to 76–80% (Lister and Murant 1967). For more details of ToRSV, refer to Solanum lycopersicum.

Tomato severe rugose virus Taxonomic position Genus: Begomovirus

(ToSRV)

Family: Geminiviridae

ToSRV infection in plants of Glycine max was reported from Brazil (Macedo et al. 2017). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ToSRV, refer to Solanum lycopersicum.

G Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Glycine max was reported from Korea, Iran, and Georgia (USA) (Golnaraghi et al. 2000, 2001, 2004; Nischwitz et al. 2006; Yoon et al. 2018). The virus-infected soybean plants do not exhibit any symptoms. The virus is transmitted by thrips vectors in a persistentpropagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow ring virus

(TYRV)

Synonyms Tomato yellow fruit ring virus (TYFRV) Taxonomic position TYRV is a tentative member of the genus Orthotospovirus and family Tospoviridae TYFRV infection in plants of Glycine max was reported from Iran (Shahraeen and Ghotbi 2003; Golnaraghi et al. 2007). The virus-infected soybean plants exhibit symptoms of leaf chlorosis, mosaic, mottling, yellowing, stunting, and vein or top necrosis. The virus is transmitted by the thrips vector, Thrips tabaci, in a persistent-propagative manner, and is also transmissible by mechanical sapinoculation. For more details of TYRV, refer to Solanum lycopersicum.

Tomato yellow spot virus Taxonomic position Genus: Begomovirus

(ToYSV)

Family: Geminiviridae

1096

Glycine max (Soybean)

ToYSV naturally infects plants of Glycine max in northwestern Argentina (Rodriguez-Pardina et al. 2011). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ToYSV, refer to Solanum lycopersicum.

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Glycine soja (Wild soybean)

Yasmin T, Nelson BD, Hobbs HA, McCoppin NK, Lambert KN, Domier LL (2017) Molecular characterization of a new soybean-infecting member of the genus Nepovirus identified by high-throughput sequencing. Arch Virol 162:1089–1092 Yi W-X, Chen S-S, Yang CY (2011) One-tube real-time PCR assay for simultaneous detection of Bean pod mottle virus and Tobacco ring spot virus in soybean seeds. Acta Phytopathol Sin 41:85–92 Yoon YN, Jo Y, Cho WK, Choi H, Jang Y, Lee YH, Bae JY, Lee BC (2018) First report of Tomato spotted wilt virus infecting soybean in Korea. Plant Dis 102:461 Zanardo LG, Carvalho CM (2017) Cowpea mild mottle virus (Carlavirus, Betaflexiviridae): a review. Trop Plant Pathol 42:417–430 Zanardo LG, Silva FN, Lima AT, Milanesi DF, Castilho-Urquiza GP, Almeida AM, Zerbini FM, Carvalho CM (2014) Molecular variability of cowpea mild mottle virus infecting soybean in Brazil. Arch Virol 159:727–737 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zettler FW, Stansly PA, Elliott MS, Fabiani AE, Scott HA, Morales FJ, Niessen AI (1989) Report of Bean pod mottle virus in South America. Plant Dis 73:518 Zhang H, Cui X, Chen X, Zhi H, Zhang S, Zhao L (2012) Determination of the complete genomic sequence and molecular biological analysis of Soybean mosaic virus. Can J Plant Pathol 34(2):288–297 Zhao F, Hwang US, Lim S, Yoo RH, Igori D, Lee SH, Lim HS, Moon JS (2015) Complete genome sequence and construction of infectious full-length cDNA clones of tobacco ringspot Nepovirus, a viral pathogen causing bud blight in soybean. Virus Genes 51(1):163–166 Zheng C, Chang R, Qiu L (2000) Progress on the study of the disease Soybean mosaic virus. Acta Phytopathol Sin 30:97–105 Zhou J (2012) Characterization and epidemiology of Soybean vein necrosis associated virus. MS thesis, University of Arkansas, Fayetteville Zhou J, Tzanetakis IE (2013) Epidemiology of Soybean vein necrosis-associated virus. Phytopathology 103:966–971 Zhou J, Tzanetakis IE (2018) Soybean vein necrosis virus: an emerging virus in North America. Virus Genes. https://doi. org/10.1007/s11262-018-1618-4 Zhou J, Tzanetakis IE (2019) Soybean vein necrosis virus: an emerging virus in North America. Virus Genes 55(1):12–21 Zhou Y, Hou Q, Yang D (1995) Identification of soybean viruses from ten provinces of China. Soybean Sci 14:246–250 Zhou J, Kantartzi SK, Wen R-H, Newman M, Hajimorad MR, Rupe JC, Tzanetakis IE (2011) Molecular characterization of a new tospovirus infecting soybean. Virus Genes 43:289–295 Zhou GC, Wu XY, Zhang YM, Wu P, Wu XZ, Liu LW, Wang Q, Hang YY, Yang JY, Shao ZQ, Wang B, Chen JQ (2014) A genomic survey of thirty soybean-infecting bean common mosaic virus (BCMV) isolates from China pointed BCMV as a potential threat to soybean production. Virus Res 191:125–133 Zinsou VA, Afouda LAC, Zoumarou-Wallis N, Dossou L, Gomez J, Soumaila F, Afloukou F, Kotchofa R (2015) Importance of cowpea mild mottle virus on soybean (Glycine max) in Benin and effect of planting date on soybean (G. max) virus level in northern Benin. Crop Protect 72:139–143

Glycine soja (Wild soybean) Family: Rosaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Weed host

(CMV)

Family: Bromoviridae

CMV infection in plants of Glycine soja was reported from Korea (Phan et al. 2014). The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of CMV, refer to Cucumis sativus.

Glycine tomentella (Wooly glycine)

Soybean mosaic virus Taxonomic position Genus: Potyvirus

1105

(SMV)

Family: Potyviridae

SMV-infected plants of Glycine soja were reported from China (Chen et al. 2017). The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of SMV, refer to Glycine max.

References Chen YX, Wu M, Ma FF, Chen JQ, Wang B (2017) Complete nucleotide sequences of seven soybean mosaic viruses (SMV), isolated from wild soybeans (Glycine soja) in China. Arch Virol 162:901–904 Phan MSV, Seo J-K, Choi H-S, Lee S-H, Kim K-H (2014) Molecular characterization of an isolate of Cucumber mosaic virus from Glycine soja by generating its infectious full-genome cDNA clones. Plant Pathol J 30:159–167

Glycine tomentella (Wooly glycine) Family: Fabaceae

Glycine mottle virus

Weed host

(GMoV)

Taxonomic position GMoV is a tentative member of the genus Carmovirus. Geographical distribution GMoV was first reported in plants of Glycine tomentella from Herberton, Queensland, Australia, by Behncken and Dale (1984). Symptoms and host(s) The virus-infected wooly glycine plants exhibit systemic mottle symptoms. Transmission The virus is transmissible by mechanical sap-inoculation. Virion properties and genome The virions are isometric, non-enveloped, and 30 nm in diameter. The genome consists of a singlestranded RNA of 4.2 kb.

References Behncken GM, Dale JL (1984) Glycine mottle virus, a possible member of the tombusvirus group. Intervirology 21:159

G

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Glycosmis pentaphylla (Orange berry)

Glycosmis pentaphylla (Orange berry) Family: Rutaceae

Citrus leprosis virus C Taxonomic position Genus: Cilevirus

Edible fruit

(CiLV-C)

Family: Kitaviridae

CiLV-C infection in plants of Glycosmis pentaphylla was reported from Brazil (Freitas-Astua et al. 2009). The virus-infected orange berry plants exhibit conspicuous dark spots, with a darker center and irregular borders surrounded by small chlorotic halo symptoms. The virus is transmitted by the mite vector, Brevipalpus phoenicis, and is also transmissible by mechanical sap-inoculation. For more details of CiLV-C, refer to Citrus spp.

References Freitas-Astua J, Fadel AL, Antonioli-Luizon R, Bastianel M, Novelli VM, Kitajima EW, Machado MA (2009) The remote citroid fruit tree Glycosmis pentaphylla is a host of Citrus leprosis virus C and exhibits novel leprosis symptoms. J Plant Pathol 91:499

Gnaphalium purpureum (Spoon-leaf purple everlasting) Family: Asteraceae

Weed host

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Gnaphalium purpureum was reported from the USA (Groves et al. 2002). The virus-infected Gnaphalium purpureum plants exhibit yellow spots or ringspots on the leaves. The virus is transmitted by the thrips vector, Frankliniella fusca, in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Groves RL, Walgenbach JF, Mayor JW, Kennedy GG (2002) The role of weed hosts and Tobacco thrips, Frankliniella fusca, in the epidemiology of Tomato spotted wilt virus. Plant Dis 86:573–582

Gomphrena globosa (Globe amaranth)

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Gompholobium preissii (Glory peas) Family: Fabaceae

Weed host

Gompholobium virus A Taxonomic position The virus is a tentative member of the genus Unassigned and family Tombusviridae. Geographical distribution The virus infection in plants of Gompholobium preissii was reported from Australia (Koh et al. 2016). Symptoms and host(s) The virus-infected glory pea plants do not exhibit any symptoms. Transmission The virus is mechanically sap-transmissible. Virion properties and genome The genome is 3935 nt (NC_030742) (Koh et al. 2016).

References Koh SH, Ong JW, Admiraal R, Sivasithamparam K, Jones MG, Wylie SJ (2016) A novel member of the Tombusviridae from a wild legume, Gompholobium preissii. Arch Virol 161:2893–2898

Gomphrena globosa (Globe amaranth) Family: Amaranthaceae

Beet mosaic virus Taxonomic position Genus: Potyvirus

Ornamental

(BtMV)

Family: Potyviridae

BtMV infection in plants of Gomphrena globosa was reported from Italy (Martelli and Russo 1969). The virus-infected globe amaranth plants exhibit chlorotic local lesion symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation, and by grafting. The virus is not transmissible by contact between plants. For more details of BtMV, refer to Beta vulgaris.

G

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Gomphrena virus

Gomphrena globosa (Globe amaranth)

(GoV)

Taxonomic position GoV is a tentative member of the family Rhabdoviridae.

Geographical distribution GoV infection in plants of Gomphrena globosa was reported from Brazil (Kitajima and Costa 1966).

Symptoms and host(s) The virus-infected globe amaranth plants exhibit local lesions.

Transmission There is no vector identified, but there are aphids suspected.

Virion properties and genome The virions are bacilliform, or bullet-shaped, measuring 230–250 nm in length and 75 nm in diameter. The genome is not segmented and contains a single molecule of linear single-stranded RNA (Kitajima and Costa 1966), presumed to be of negative sense.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Gomphrena globosa was reported from Eastern Washington (Kaiser et al. 1982). The virus is transmitted by the thrips vectors; the virus present in/on pollen entering the host through injuries caused by thrips while feeding. The virus is mechanically sap-transmissible, but is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Gomphrena globosa was reported from Pennsylvania (USA) (Hausbeck et al. 1992). The virus-infected globe amaranth plants exhibit necrosis, ringspots, and chlorotic symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Gossypium spp. (Cotton)

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References Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among green house ornamentals in Pennsylvania. Plant Dis 76:795–800 Kaiser WJ, Wyatt SD, Pesho GR (1982) Natural hosts and vectors of tobacco streak virus in eastern Washington. Phytopathology 72:1508–1512 Kitajima EW, Costa AS (1966) Morphology and developmental stages of Gomphrena virus. Virology 29:523 Martelli GP, Russo M (1969) Nuclear changes in mesophyll cells of Gomphrena globosa L. associated with infection by beet mosaic virus. Virology 38:297–308

Gossypium spp. (Cotton) Family: Malvaceae

Fiber crop

Chickpea chlorotic dwarf virus Taxonomic position Genus: Mastrevirus

G (CpCDV)

Family: Geminiviridae

CpCDV infection in plants of Gossypium spp. was reported from Punjab province, Pakistan (Manzoor et al. 2014). The virus-infected cotton plants exhibit symptoms consisting of leaf curling, vein darkening, vein thickening, and enations that sometimes develop into cup-shaped, leaflike structures on veins on the underside of the leaves. The virus is transmitted by the leafhopper vector, Orosius albicinctus, in a persistent, circulative, and non-propagative manner, and infects a number of economically important crops. The virus is not transmissible by mechanical inoculation. For more details of CpCDV, refer to Cicer arietinum.

Cotton chlorotic spot virus Taxonomic position Genus: Begomovirus

(CoChSpV)

Family: Geminiviridae

Geographical distribution CoChSpV infection in plants of Gossypium hirsutum x Gossypium mustelinum was reported from Brazil (de Almeida et al. 2013). Symptoms and host(s) The virus-infected cotton plants exhibit chlorotic spotting symptoms. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single-coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A

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Gossypium spp. (Cotton)

contains 2670 nt (KF358470) and DNA-B of 2650 nt (KF358471) (Briddon 2001; de Almeida et al. 2013; Brown et al. 2015; Zerbini et al. 2017).

Cotton leaf crumple virus Taxonomic position Genus: Begomovirus

(CLCrV)

Family: Geminiviridae

Geographical distribution CLCrV infection in plants of Gossypium spp. was reported from southwestern Arizona, Guatemala, Texas, California, Mexico, and India (Dickson et al. 1954; Allen et al. 1960; Mali 1977; Brown 1992; Idris and Brown 2004). Earlier the virus is known as “acromania” or “crazy top” (Cook 1924). Symptoms and host(s) The virus has only been detected in Gossypium hirsutum, bean (Phaseolus vulgaris), Abutilon theophrasti, and cheeseweed (Malva parviflora). In Gossypium hirsutum, CLCrV induces crumpling, foliar discoloration, and an overall stunting of plants when infection occurs at early growth stages. The most pronounced symptoms are usually observed on immature leaves. Minute outgrowths are often observed on flowers of infected plants and bolls can be misshapen (Brown 1992). The symptom on the other hosts was not noted. Transmission The virus is transmitted by the whitefly vector of begomoviruses, Bemisia tabaci. The minimum acquisition access period was found to be 1 h and the minimum inoculation access period 4 h. The latent period was estimated at 24–28 h. The mechanism of transmission has not been determined but is likely circulative and non-propagative, in common with other begomoviruses. The virus lost by the vector when it molts does not multiply in the vector and not transmitted congenitally to the progeny of the vector. CLCrV has a narrow host range and was experimentally transmitted by whitefly inoculation only to a few host species belonging to the Malvaceae and Fabaceae: Gossypium barbadense, G. hirsutum, G. thruberi, Abutilon theophrasti, Alcea rosea, Hibiscus cameronii, H. cannabinus, H. cameronii, H. palustris, H. diversifolius, H. palustris, H. sabdariffa, Malva parviflora, Castanospermum australe, Glycine max, Phaseolus aureus, P. angularis and P. acutifolius on which foliar blistering, mosaic, and malformation symptoms are produced (Brown et al. 1986; Brown and Nelson 1987). The virus is not transmitted through seed or sap. The virus is not transmissible by contact between plants and not transmitted by pollen. The virus is transmissible by grafting (Brown and Nelson 1987). Virion properties and genome CLCrV virions are typically geminate and ~20  30 nm in size with a paired quasi-icosahedral morphology (Brown and Nelson 1984). CLCrV is typical of the majority of begomoviruses native to the New World with a bipartite genome consisting of two circular, single-stranded DNA components. DNA-A contains 2630 nt (AF480940 = NC_004580; AY742220) and DNA-B of 2550 nt (AF480941 = NC_004581; AY742221) (Briddon 2001; Brown and Idris 2002; Idris and Brown 2004; Brown et al. 2015; Zerbini et al. 2017). Typical of all bipartite begomoviruses native to the New World, the DNA-A component of CLCrV encodes five genes, one in the virion-sense and four in the complementary-sense, whereas the

Gossypium spp. (Cotton)

1111

DNA-B encodes one gene in each orientation. The expression and function of these genes has not been investigated for CLCrV.

Cotton leaf curl Alabad virus Taxonomic position Genus: Begomovirus

(CLCuAlV)

Family: Geminiviridae

Geographical distribution CLCuAlV infection in plants of Gossypium spp. was reported from Alabad (Pakistan) (Zhou et al. 1998; Nawaz-ul-Rehman et al. 2012). Symptoms and host(s) The virus-infected cotton plants exhibit typical cotton leaf curl disease symptoms (leaf curling, vein swelling, enations, leaflike enations, vein darkening, and stunting). Transmission The virus is transmitted by the whitefly vector Bemisia tabaci. The mechanism of transmission has not been determined but is likely circulative and non-propagative, in common with other begomoviruses. Virion properties and genome The structure of the virions of CLCuBaV has not been investigated. In common with all geminiviruses, the virions of CLCuBaV are likely geminate (twinned quasi-icosahedra). CLCuAlV is unusual in that some isolates appear to have monopartite genomes (those isolated from Gossypium hirsutum and okra (AJ002455 = NC_004582; AJ002452, FJ218485, GU112004, GU112081)) and some appear to have bipartite genomes (those isolated from the exotic cotton species Gossypium punctatum, Gossypium davidsonii, and Gossypium lobatum (DNA-A: EU365617, EU384575, FJ210467 and DNA-B: EU384577, EU384578, FJ218488)). CLCuAlV is a typical Old World begomovirus. The genome consists of either a single circular molecule of a single-stranded DNA of ~2750 nt or two circular molecules of a single-stranded DNA each of ~2750 nt. The genome or DNA-A component encodes the six genes typically encoded by monopartite begomoviruses originating from the Old World. The DNA-B component (associated with some isolates) encodes one gene in each orientation. The expression and function of the genes have not been investigated. Although the characterized monopartite isolates of CLCuAlV were not shown to be associated with a betasatellite, it is likely that a betasatellite is associated with the virus, in common with other monopartite begomoviruses that infect cotton and okra. CLCuAlV is a part of the virus-betasatellite complex that has been causing cotton leaf curl disease in Pakistan and northwestern India since the early 1990s (Briddon 2001; Zhou et al. 1998; Nawaz-ul-Rehman et al. 2012; Brown et al. 2015; Zerbini et al. 2017).

Cotton leaf curl Bangalore virus Taxonomic position Genus: Begomovirus

(CLCuBaV)

Family: Geminiviridae

G

1112

Gossypium spp. (Cotton)

Geographical distribution CLCuBaV infection in plants of Gossypium barbadense was reported from Bangalore (Southern India) (Chowda Reddy et al. 2005). Symptoms and host(s) The virus-infected cotton plants exhibit typical cotton leaf curl disease symptoms (leaf curling, enations, vein swelling). Transmission The virus is transmitted by the whitefly vector Bemisia tabaci. The minimum acquisition access period was determined to be 15 min and the minimum inoculation access period 10 min. The mechanism of transmission has not been determined but is likely circulative and non-propagative, in common with other begomoviruses. Virion properties and genome The structure of the virions of CLCuBaV has not been investigated. In common with all geminiviruses, the virions of CLCuBaV are likely geminate (twinned quasi-icosahedra). CLCuBaV is a typical Old World monopartite begomovirus. Only two isolates of the virus have been characterized, and the genome consists of a single circular molecule of a single-stranded DNA of ~2751 nt (AY705380 = NC_007290; GU112003) (Briddon 2001; Chowda Reddy et al. 2005; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes encode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated. Although the characterized isolates of CLCuBaV were not shown to be associated with a betasatellite, it is likely that a betasatellite is associated with the virus, in common with other monopartite begomoviruses that infect cotton and okra. CLCuBaV is not a part of the virusbetasatellite complex that has been causing cotton leaf curl disease in Pakistan and northwestern India since the early 1990s.

Cotton leaf curl Gezira virus Taxonomic position Genus: Begomovirus

(CLCuGeV)

Family: Geminiviridae

Geographical distribution CLCuGeV infection in plants of Gossypium spp. was reported from Gezira (Sudan) and Pakistan (Zhou et al. 1998; Idris and Brown 2000; Tahir et al. 2011). Symptoms and host(s) The virus-infected cotton plants exhibit typical cotton leaf curl symptoms (leaf curling, vein swelling, vein darkening, and enations). The virus has been detected in Hibiscus esculentus, Sida alba and Carica papaya. Transmission CLCuGeV has been shown to be transmitted by the whitefly vector Bemisia tabaci (Idris and Brown 2000). It is likely that, in common with other begomoviruses, the mechanism of transmission of CLCuGeV by B. tabaci is in a circulative, non-propagative manner.

Gossypium spp. (Cotton)

1113

Experimentally, a CLCuGeV isolate from Mali, isolated from okra, was shown to infect okra, but not cotton (Gossypium hirsutum), by Agrobacterium-mediated inoculation together with the cognate betasatellite, inducing typical leaf curling and thickening of small veins. Virion properties and genome The structure of the virions of CLCuGeV has not been investigated. In common with all geminiviruses, the virions of CLCuGeV are likely geminate (twinned quasi-icosahedra). CLCuGeV is a typical Old World monopartite begomovirus. The genome of CLCuGeV consists of a single circular molecule of a single-stranded DNA of 2764 nt (AF260241 = NC_002510; AY036008, HG969203) (Briddon 2001; Idris et al. 2005; Tiendrebeogo et al. 2010; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes of CLCuGeVencode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes of CLCuGeV have not been investigated. Cotton leaf curl Gezira betasatellite DNA molecule is associated with CLCuGeV, and consists of 1349 nt (DQ644564) (Zhou 2013).

Cotton leaf curl Kokhran virus

(CLCuKoV)

Synonyms Cotton leaf curl Burewala virus (CLCuBuV) Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution CLCuKoV infection in plants of Gossypium hirsutum was reported from Faisalabad (Pakistan) and India (Mahmood et al. 2003; Mansoor et al. 2003; Kirthi et al. 2004; Kumar et al. 2011; Zaffalon et al. 2011; Hina et al. 2012; Shuja et al. 2014). Symptoms and host(s) The virus-infected cotton plants exhibit leaf curling symptoms. The virus has been isolated from cotton (mainly Gossypium hirsutum and G. barbadense but also other tetraploid Gossypium species.), okra (Abelmoschus esculentus), tomato (Solanum lycopersicum), Cyamopsis tetragonoloba, Luffa cylindrica, and Xanthium strumarium. Transmission Although the transmission of CLCuKoV has not been investigated, the virus complex (virus not identified) causing CLCuD has been transmitted. The complex was transmitted by the whitefly vector, Bemisia tabaci, in a persistent manner (Mansoor et al. 1993; Sharma and Rishi 2003). The virus was transmitted from cotton to cotton by adult B. tabaci after an acquisition access period of 1 h, could be inoculated in 5 min, had a minimum latent period of 8 h, and was retained by viruliferous insects for up to 9 days. Female B. tabaci transmitted more frequently than males. The complex is graft-transmissible but not mechanically transmissible. Experimentally, the virus has been introduced into cotton (Gossypium hirsutum) Nicotiana benthamiana and tobacco (N. tabacum) by both Agrobacterium-mediated inoculation and biolistic inoculation of the virus genome (Mansoor et al. 2003). Care must be taken in interpreting both the host range of and symptoms induced byCLCuKoV since the host range and symptoms of this virus are, in many cases, determined by the betasatellite with which it is associated.

G

1114

Gossypium spp. (Cotton)

Virion properties and genome The structure of the virions of CLCuKoV has not been investigated. In common with all geminiviruses, the virions of CLCuKoV are likely geminate (twinned quasi-icosahedra). CLCuKoV is a typical Old World monopartite begomovirus. The genome of CLCuKoV consists of a single circular molecule of a single-stranded DNA of 2748 nt (AJ002449 = NC_004583; AJ002448) (Zhou et al. 1998; Briddon 2001; Mansoor et al. 2003; Amrao et al. 2010; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes of CLCuKoV encode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated. CLCuKoV has been shown to be associated with a betasatellite.

Cotton leaf curl Multan virus

(CLCuMuV)

Synonyms Cotton leaf curl Rajasthan virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution CLCuMuV infection in plants of Gossypium spp. was reported from Pakistan and China (Zhou et al. 1998; Kirthi et al. 2004; Radhakrishnan et al. 2004a, b; Du et al. 2015; Cai et al. 2010; Sattar et al. 2013; Gu et al. 2015). Symptoms and host(s) CLCuMuV was shown to poorly infect cotton, in the absence of CLCuMuB, and to induce atypical (mild) symptoms in cotton (Briddon et al. 2000). In the presence of CLCuMuB, CLCuMuV induced typical CLCuD symptoms that include leaf curling, vein swelling, vein darkening, enations, and leaflike enations (Briddon et al. 2001; Qazi et al. 2007). CLCuMuV has been mainly isolated from Gossypium hirsutum and G. barbadense and also other tetraploid Gossypium species, okra (Abelmoschus esculentus), tomato (Solanum lycopersicum), rose mallow (Hibiscus rosa-sinensis), Kenaf (Hibiscus cannabinus), papaya (Carica papaya), Malvaviscus penduliflorus, and Digera arvensis (Mubin et al. 2009). Transmission Although the transmission of CLCuMuV has not been investigated, the virus complex (virus not identified) causing CLCuD has been transmitted. The complex was transmitted by the whitefly vector Bemisia tabaci in a persistent manner (Sharma and Rishi 2003; Mansoor et al. 1993). The virus was transmitted from cotton to cotton by adult B. tabaci after an acquisition access period of 1 h, could be inoculated in 5 min, had a minimum latent period of 8 h, and was retained by viruliferous insects for up to 9 days. Female B. tabaci transmitted more frequently than males. The complex is graft-transmissible but not mechanically transmissible. Experimentally, the virus has been transmissible to cotton (Gossypium hirsutum) Nicotiana benthamiana and tobacco (N. tabacum) by both Agrobacterium-mediated inoculation and biolistic inoculation of the virus genome (Briddon et al. 2001; Mansoor et al. 2003).

Gossypium spp. (Cotton)

1115

Virion properties and genome The structure of the virions of CLCuMuV has not been investigated. In common with all geminiviruses, the virions of CLCuMuV are likely geminate (twinned quasi-icosahedra). CLCuMuV is a typical Old World monopartite begomovirus. The genome of CLCuMuV consists of a single circular molecule of a single-stranded DNA of 2751 nt (AJ002458, AJ002447, AJ496287, X98995) (Zhou et al. 1998; Briddon et al. 2000; Briddon 2001; Mansoor et al. 2003; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes of CLCuMuV encode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated. Cotton leaf curl Multan betasatellite DNA molecule is associated with CLCuMuV, and consists of 1349 nt (AJ298903) (Briddon et al. 2001, 2003; Amin et al. 2006; Nawaz-Ul-Rehman et al. 2010; Yang et al. 2011; Zhou 2013).

Cotton leafroll dwarf virus Taxonomic position Genus: Polerovirus

(CLRDV)

Family: Luteoviridae

Geographical distribution CLRDV is commonly known as the agent of cotton blue disease. CLRDV infection in plants of Gossypium spp. was reported from Argentina, South America, America, Africa, Thailand, and Asia (Distefano et al. 2010; Mukherjee et al. 2012; Delfosse et al. 2013; Sharman et al. 2015a; Ray et al. 2016; Agrofoglio et al. 2017). Symptoms and host(s) The virus-infected cotton plants exhibit stunting, leaf rolling, intense green foliage, vein yellowing, brittleness of leaves, and reduced flower and boll size, sometimes resulting in sterility of plants. Transmission The virus is transmitted by the aphid vector, Aphis gossypii, in a circulative and non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. Virion properties and genome The virions are 25–30 nm in diameter, hexagonal in outline, and have no envelope. The genome is a positive-sense, single-stranded RNA of 5866 nt (GU167940). The 30 terminus has neither a poly (A) tract nor a tRNA-like structure. The 50 terminus has a genome-linked protein (VPg) of about 17 kDa. The genome contains six open reading frames (ORFs) (Correa et al. 2005; Distefano et al. 2010; Cascardo et al. 2015).

Cotton yellow mosaic virus (CYMV) Taxonomic position Genus: Begomovirus Family: Geminiviridae Geographical distribution CYMV infection in plants of Gossypium raimondii was reported from Benin, West Africa (Leke et al. 2016).

G

1116

Gossypium spp. (Cotton)

Symptoms and host(s) The virus-infected cotton plants exhibit light yellow mosaic symptoms. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm, with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2766 nt (KT454834 = NC_030310; KU683748) and DNA-B of 2716 nt (KT454835 = NC_030311) (Briddon 2001; Brown et al. 2015; Leke et al. 2016; Zerbini et al. 2017).

Groundnut bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(GBNV)

Family: Tospoviridae

GBNV infection in plants of Gossypium hirsutum was reported from India (Jain et al. 2004). The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a number hosts. For more details of GBNV, refer to Arachis hypogaea.

Okra enation leaf curl virus Taxonomic position Genus: Begomovirus

(OELCuV)

Family: Geminiviridae

OELCuV infection in plants of Gossypium hirsutum was reported from Pakistan (Hameed et al. 2014). The virus-infected cotton plants exhibit symptoms of vein thickening, upward or downward leaf curling, and foliar enations. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of OELCuV, refer to Abelmoschus esculentus.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Gossypium spp. was reported from Australia, India, and Pakistan (Bhat et al. 2002; Ahmed et al. 2003; Waqar et al. 2003; Sharma 2006; Sharman et al. 2008, 2015b; Prasada Rao et al. 2009; Jagtap et al. 2012a; Arutselvan 2013; Rageshwari et al. 2016; Vinodkumar et al. 2017). The virus-infected cotton plants show clear chlorosis and mosaic symptoms on major leaves of plants. Initial infection of leaves results in chlorotic lesions which later turn into necrotic, purplish brown spots and rings; the leaf size is reduced, and there are blisters along the veins and mosaic patterns between secondary veins. Diseased cotton plants are slightly stunted and produce more axillary twigs than healthy plants. Even necrosis of the buds and drying up of young bolls was also observed.

Gossypium spp. (Cotton)

1117

The virus is transmitted by the thrips vectors (Frankliniella schultzei, Thrips palmi, and Thrips tabaci); the virus present in/on pollen entering the host through injuries caused by thrips while feeding (Jagtap et al. 2012b). The virus is mechanically sap-transmissible to Malvaceae, Chenopodiaceae, Compositae, Leguminosae, and Solanaceae members. The virus is pollen transmitted. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato leaf curl Gujarat virus Taxonomic position Genus: Begomovirus

(ToLCGuV)

Family: Geminiviridae

ToLCGuV infection in plants of Gossypium hirsutum was reported from Pakistan and northwestern India (Zaidi et al. 2015). The virus-infected cotton plants exhibit leaf curling and vein thickening symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, nonpropagative manner, and also by grafting. For more details of ToLCGuV, refer to Solanum lycopersicum.

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Gossypium hirsutum was reported from Pakistan (Zaidi et al. 2016). The virus-infected cotton plants exhibit leaf curling, vein thickening, vein yellowing, enations, and stunted growth. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, nonpropagative manner. The virus is graft-transmissible. For more details of ToLCNDV, refer to Solanum lycopersicum.

References Agrofoglio YC, Delfosse VC, Casse MF, Hopp HE, Kresic IB, Distefano AJ (2017) Identification of a new cotton disease caused by an a typical Cotton leaf roll dwarf virus in Argentina. Phytopathology 107:369–376 Ahmed W, Butt TB, Ihsan J, Rehman A (2003) Natural occurrence of Tobacco streak virus in cotton in Pakistan and screening for its resistant sources. Pak J Bot 35:401–408 Allen RM, Tucker H, Nelson RA (1960) Leaf crumple disease of cotton in Arizona. Plant Dis Reptr 44:246–250 de Almeida MM, Jain S, Barroso PA, Hoffmann LV, de Lucena MG, de Resende RO, Inoue-Nagata AK (2013) Complete sequence of a new bipartite begomovirus infecting cotton plants in Brazil. Genome Announc 1(6):e00661–e00613 Amin I, Mansoor S, Amrao L, Hussain M, Irum S, Zafar Y, Bull SE, Briddon RW (2006) Mobilisation into cotton and spread of a recombinant cotton leaf curl disease satellite. Arch Virol 151(10):2055–2065 Amrao L, Amin I, Shahid MS, Briddon RW, Mansoor S (2010) Cotton leaf curl disease in resistant cotton is associated with a single begomovirus that lacks an intact transcriptional activator protein. Virus Res 152(1–2):153–163 Arutselvan R (2013) Studies on Tobacco streak virus infecting cotton in Coimbatore and Erode district in Tamil Nadu. M. Sc.(ag.) thesis, Tamil Nadu Agricultural University Bhat AI, Jain RK, Chaudhury V, Krishnareddy M, Ramaiah K, Chattannavar SN, Varma A (2002) Sequence conservation in the coat protein gene of Tobacco streak virus isolates causing necrosis disease in cotton, mungbean, sunflower, and sunn-hemp in India. Indian J Biotechnol 1:350–356 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer index of viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55

G

1118

Gossypium spp. (Cotton)

Briddon RW, Mansoor S, Bedford ID, Pinner MS, Markham PG (2000) Clones of cotton leaf curl geminivirus induce symptoms a typical of cotton leaf curl disease. Virus Genes 20(1):19–26 Briddon RW, Mansoor S, Bedford ID, Pinner MS, Saunders K, Stanley J, Zafar Y, Malik KA, Markham PG (2001) Identification of dna components required for induction of cotton leaf curl disease. Virology 285(2):234–243 Briddon RW, Bull SE, Amin I, Idris AM, Mansoor S, Bedford ID, Dhawan P, Rishi N, Siwatch SS, Abdel-Salam AM, Brown JK, Zafar Y, Markham PG (2003) Diversity of DNA b: a satellite molecule associated with some monopartite begomoviruses. Virology 312(1):106–121 Brown JK (1992) Virus diseases of cotton. In: Hillocks RJ (ed) Cotton diseases. Common Wealth Agricultural Bureaux International, Oxon, p 415 Brown JK, Idris AM (2002) Cotton leaf crumple virus is the first member of a previously undiscovered phylogenetic group of Western Hemisphere begomoviruses. Phytopathology 92:S38 Brown JK, Nelson MR (1984) Geminate particles associated with cotton leaf crumple disease in Arizona. Phytopathology 74(8):987–990 Brown JK, Nelson MR (1987) Host range and vector relationships of Cotton leaf crumple virus. Plant Dis 71:522–524 Brown JK, Nelson MR, Lambe RC (1986) Cotton leaf crumple virus transmitted from naturally infected bean from Mexico. Plant Dis 70:981 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Cai JH, Xie K, Lin L, Qin BX, Chen BS, Meng JR, Liu YL (2010) Cotton leaf curl Multan virus newly reported to be associated with cotton leaf curl disease in China. Plant Pathol 59:794–795 Cascardo RS, Arantes ILG, Silva TF, Sachetto-Martins G, Vaslin MFS, Correa RL (2015) Function and diversity of P0 proteins among cotton leafroll dwarf virus isolates. Virol J 12:123 Chowda Reddy RV, Muniyappa V, Colvin J, Seal S (2005) A new begomovirus isolated from Gossypium barbadense in Southern India. Plant Pathol 54:570 Cook OF (1924) Acromania or crazy top, a growth disorder of cotton. J Agric Res 28:803–828 Correa RL, Silva TF, Simoes-Araújo JL, Barroso PAV, Vidal MS, Vaslin MFS (2005) Molecular characterization of a virus from the family Luteoviridae associated with cotton blue disease. Arch Virol 150:1357–1367 Delfosse VC, Casse MF, Agrofoglio YC, Kresic IB, Hopp HE, Ziegler-Graff V, Distefano AJ (2013) Agroinoculation of a full-length cDNA clone of cotton leafroll dwarf virus (CLRDV) results in systemic infection in cotton and the model plant Nicotiana benthamiana. Virus Res 175:64–70 Dickson RC, Johnson MM, Laird EF (1954) Leaf crumple, a virus disease of cotton. Phytopathology 44:479–480 Distefano AJ, Bonacic Kresic I, Hopp HE (2010) The complete genome sequence of a virus associated with cotton blue disease, cotton leafroll dwarf virus, confirms that it is a new member of the genus Polerovirus. Arch Virol 155(11):1849–1854 Du Z, Tang Y, He Z, She X (2015) High genetic homogeneity points to a single introduction event responsible for invasion of Cotton leaf curl Multan virus and its associated betasatellite into China. Virol J 12(1):163 Gu Z-H, Hu G-J, Xie Y, Zhou X-P (2015) Construction and pathogenicity evaluation of infectious clones of Cotton leaf curl Multan virus infecting Hibiscus rosasinensis and Gossypium hirsutum [J]. Acta Phytopathol Sin 45(2):198–204 Hameed U, Zia-Ur-Rehman M, Harrmann HW, Heider MS, Brown JK (2014) First report of Okra enation leaf curl virus and associated Cotton leaf curl Multan betasatellite and Cotton leaf curl Multan alphasalite infecting cotton in Pakistan: a new member of the Cotton leaf curl disease complex. Plant Dis 98:1447–1448 Hina S, Arshad Javed M, Haider S, Saleem M (2012) Isolation and sequence analysis of cotton infecting begomoviruses. Pak J Bot 44:223–230 Idris AM, Brown JK (2000) Identification of a new, monopartite begomovirus associated with leaf curl disease of cotton in Gezira, Sudan. Plant Dis 84:809 Idris AM, Brown JK (2004) Cotton leaf crumple virus is a distinct Western Hemisphere begomovirus species with complex evolutionary relationships indicative of recombination and reassortment. Phytopathology 94:1068–1074 Idris AM, Briddon RW, Bull SE, Brown JK (2005) Cotton leaf curl Gezira virus-satellite DNAs represent a divergent, geographically isolated Nile Basin lineage: predictive identification of a satDNA REP-binding motif. Virus Res 109(1):19–32 Jagtap GP, Jadhav TH, Utpal D (2012a) Occurrence, distribution and survey of Tobacco streak virus (TSV) of cotton. J Plant Dis Sci 7:251–253 Jagtap GP, Jadhav TH, Utpal D (2012b) Host range and transmission of Tobacco streak virus (TSV) causing cotton mosaic disease. Sci J Vet Adv 1(1):22–27 Jain RK, Sundeep B, Ramiah M (2004) Natural infection of Groundnut bud necrosis virus in cotton and sem (Abstract). In: National symposium on molecular diagnostics for the management of viral diseases. October, 14–16, 2004. Indian Agricultural Research Institute, New Delhi.

Gossypium spp. (Cotton)

1119

Kirthi N, Priyadarshini CGP, Sharma P, Maiya P, Hemalatha V, Sivaraman P, Dhawan P, Rishi N, Savithri HS (2004) Genetic variability of begomoviruses associated cotton leaf curl disease originating from India. Arch Virol 149:2047–2057 Kumar A, Snehi SK, Raj SK, Kumar J, Khan JA (2011) Association of Cotton leaf curl Burewala virus and its satellite molecules with leaf distortion symptoms of cotton in India. New Dis Rep 24:18 Leke WN, Khatabi B, Mignouna DB, Brown JK, Fondong VN (2016) Complete genome sequence of a new bipartite Begomovirus infecting cotton in the Republic of Benin in West Africa. Arch Virol 161:2329–2333 Mahmood T, Arshad M, Gill MI, Mahmood HT, Tahir M, Hussain S (2003) Burewala strain of Cotton leaf curl virus: a threat to CLCuV cotton resistant varieties. Asian J Plant Sci 2:968–970 Mali VR (1977) Cotton leaf crumple virus disease – a new record of India. Indian Phytopathol 30:326–329 Mansoor S, Bedford I, Pinner MS, Stanley J, Markham PG (1993) A whitefly transmitted geminivirus associated with cotton leaf curl disease in Pakistan. Pak J Bot 25:105–107 Mansoor S, Briddon RW, Bull SE, Bedford ID, Bashir A, Hussain M, Saeed M, Zafar Y, Malik KA, Fauquet C, Markham PG (2003) Cotton leaf curl disease is associated with multiple monopartite begomoviruses supported by single DNA beta. Arch Virol 148(10):1969–1986 Manzoor MT, Ilyas M, Shafiq M, Saleem Haider M, Shahid AA, Briddow RW (2014) A distinct strain of chickpea chlorotic dwarf virus (genus Mastrevirus, family Geminiviridae) identified in cotton plants affected by leaf curl disease. Arch Virol 159:1217–1221 Mubin M, Briddon RW, Mansoor S (2009) Diverse and recombinant DNA betasatellites are associated with a begomovirus disease complex of Digera arvensis, a weed host. Virus Res 142(1–2):208–212 Mukherjee AK, Chahande PR, Meshram MK, Kranthi KR (2012) First report of Polerovirus of the family Luteoviridae infecting cotton in India. New Dis Rep 25:22 Nawaz-Ul-Rehman MS, Nahid N, Mansoor S, Briddon RW, Fauquet CM (2010) Post-transcriptional gene silencing suppressor activity of two non-pathogenic alphasatellites associated with a Begomovirus. Virology 405(2):300–308 Nawaz-ul-Rehman MS, Briddon RW, Fauquet CM (2012) A melting pot of Old World begomoviruses and their satellites infecting a collection of Gossypium species in Pakistan. PLoS One 7(8):e40050 Prasada Rao RDVJ, Jyotsna MK, Reddy AS, Varaprasad KS, Nigam SN, Lava Kumar P (2009) Non-systemic infection of Tobacco streak virus on cotton in Warangal district, Andhra Pradesh. Indian J Plant Prot 37:196–198 Qazi J, Amin I, Mansoor S, Iqbal J, Briddon RW (2007) Contribution of the satellite encoded gene bC1 to cotton leaf curl disease symptoms. Virus Res 128(1–2):135–139 Radhakrishnan G, Malathi VG, Varma A (2004a) Detection of DNA-A and DNA-ß associated with cotton leaf curl and some other plant diseases caused by whitefly transmitted geminiviruses. Indian Phytopathol 57:53–60 Radhakrishnan G, Malathi VG, Varma A (2004b) Biological characterization of an isolate of Cotton leaf curl Rajasthan virus from northern India and identification of sources of resistance. Indian Phytopathol 57:174–180 Rageshwari S, Renukadevi P, Malathi VG, Nakkeeran S (2016) Occurrence, biological and serological assay of TSV infecting cotton in Tamil Nadu. J Mycol Plant Pathol 46(2):159–168 Ray JD, Sharman M, Quintao V, Rossel B, Westaway J, Gambley C (2016) Cotton leafroll dwarf virus detected in TimorLeste. Aust Plant Dis Notes 11:29 Sattar MN, Kvarnheden A, Saeed M, Briddon RW (2013) Cotton leaf curl disease – an emerging threat to cotton production worldwide. J Gen Virol 94:695–710 Sharma OP (2006) Diseases of cotton. Technical Bulletin (ICAR). pp 13–14 Sharma P, Rishi N (2003) Host range and vector relationships of Cotton leaf curl geminivirus (CLCuV) from northern India. Indian Phytopathol 56:496–499 Sharman M, Thomos JE, Persley DM (2008) First report of Tobacco streak virus in sunflower (Helianthus annuus), cotton (Gossypium hirsutum), chickpea (Cicer arietinum), mungbean (Vigna radiata) in Australia. Aust Plant Dis Notes 3:27–29 Sharman M, Lapbanjob S, Sebunruang P, Belot J-L, Galbieri R, Giband M, Suassuna N (2015a) First report of Cotton leafroll dwarf virus in Thailand using a species-specific PCR validated with isolates from Brazil. Aust Plant Dis Notes 10:24 Sharman M, Thomas JE, Persley DM (2015b) Natural host range, thrips and seed transmission of distinct Tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207 Shuja MN, Briddon RW, Tahir M (2014) Identification of a distinct strain of Cotton leaf curl Burewala virus. Arch Virol 159:2787–2790 Tahir MN, Amin I, Briddon RW, Mansoor S (2011) The merging of two dynasties – identification of an African cotton leaf curl disease-associated begomovirus with cotton in Pakistan. PLoS One 6:e20366 Tiendrebeogo F, Lefeuvre P, Hoareau M, Villemot J, Konate G, Traore AS, Barro N, Traore VS, Reynaud B, Traore O, Lett JM (2010) Molecular diversity of cotton leaf curl Gezira virus isolates and their satellite DNAs associated with okra leaf curl disease in Burkina Faso. Virol J 7:48 Vinodkumar S, Nakkeeran S, Malathi VG, Karthikeyan G, Amala Balu P, Mohankumar S, Renukadevi P (2017) Tobacco streak virus: an emerging threat to cotton cultivation in India. Phytoparasitica 45:729–743 Waqar A, Butt TB, Insan J, Rehman A (2003) Natural occurrence of Tobacco streak virus in cotton in Pakistan and screening for its resistance sources. Pak J Bot 35:401–408

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1120

Grammatophyllum spp.

Yang X, Xie Y, Raja P, Li S, Wolf JN, Shen Q, Bisaro DM, Zhou X (2011) Suppression of methylation-mediated transcriptional gene silencing by bC1-SAHH protein interaction during geminivirus-betasatellite infection. PLoS Pathog 7(10):e1002329 Zaffalon V, Mukherjee S, Reddy V, Thompson J, Tepfer M (2011) A survey of geminiviruses and associated satellite DNAs in the cotton-growing areas of northwestern India. Arch Virol 157(3):483–495 Zaidi SSA, Iqbal Z, Amin I (2015) First report of Tomato leaf curl Gujarat virus, a bipartite begomovirus on cotton showing leaf curl symptoms in Pakistan. Plant Dis 99:1655 Zaidi SSA, Shafiq M, Amin I, Scheffler BE, Scheffler JA, Briddon RW, Mansoor S (2016) Frequent occurrence of Tomato leaf curl New Delhi virus in cotton leaf curl disease affected cotton in Pakistan. PLoS One 2016 11(5):e0155520 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133 Zhou X (2013) Advances in understanding begomovirus satellites. Annu Rev Phytopathol 51:357–381 Zhou X, Liu Y, Robinson DJ, Harrison BD (1998) Four DNA-A variants among Pakistani isolates of cotton leaf curl virus and their affinities to DNA-A of geminivirus isolates from okra. J Gen Virol 79(PT 4):915–923

Grammatophyllum spp. Family: Orchidaceae

Ornamental

Cymbidium mosaic virus Taxonomic position Genus: Potexvirus

(CymMV)

Family: Alphaflexiviridae

CymMV was reported from plants of Grammatophyllum spp. in the USA (Hu et al. 1993). The virusinfected grammatophyllum plants exhibit floral and foliar necrotic symptoms. The virus is mechanically sap-transmissible and also by contact between plants. No vector transmission is reported. For more details of CymMV, refer to Cymbidium spp.

References Hu JS, Ferreira S, Wang M, Xu MQ (1993) Detection of Cymbidium mosaic virus, Odontoglossum ringspot virus, Tomato spotted wilt virus, and potyviruses infecting orchid in Hawaii. Plant Dis 77:464–468

Graptopetalum paraguayense (Ghost plant) Family: Crassulaceae

Kalanchoe latent virus Taxonomic position Genus: Carlavirus

Ornamental

(KLV)

Family: Betaflexiviridae

Griselinia lucida (Shining broadleaf)

1121

KLV infection in plants of Graptopetalum paraguayense plants reported from Italy (Sorrentino et al. 2017). The virus-infected ghost plants exhibit symptoms consisting of greenish to dark green spots visible both on old and young leaves and a mild speckling present mostly on the older leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of KLV, refer to Kalanchoe blossfeldiana.

Kalanchoe mosaic virus Taxonomic position Genus: Potyvirus

(KMV)

Family: Potyviridae

KMV infection in plants of Graptopetalum paraguayense was reported from Italy (Sorrentino et al. 2017). The virus-infected ghost plants exhibit symptoms consisting of greenish to dark green spots visible both on old and young leaves and a mild speckling present mostly on the older leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sapinoculation. For more details of KMV, refer to Kalanchoe blossfeldiana.

References Sorrentino R, Marais A, Faure C, Theil S, Alioto D, Candresse T (2017) First report of Kalanchoe mosaic virus and Kalanchoe latent virus infecting ghost plant (Graptopetalum paraguayense) in Italy. Plant Dis 101:1560

Griselinia lucida (Shining broadleaf) Family: Griseliniaceae

Tomato mosaic virus Taxonomic position Genus: Tobamovirus

Ornamental

(ToMV)

Family: Virgaviridae

ToMV infection in plants of Griselinia lucida was reported from New Zealand (Veerakone et al. 2010). The virus-infected shining broadleaf plants exhibit symptoms of chlorotic mosaic and concentric ring symptoms osn the leaves. No vector is involved in the spread of this virus. This virus is transmissible by mechanical sap-inoculation, and also by contact between plants. For more details of ToMV, refer to Solanum lycopersicum.

References Veerakone S, Lebas BSM, Tang J, Clover GRG (2010) First report of Tomato mosaic virus in Griselinia lucida, an epiphytic shrub native to New Zealand. Aust Plant Dis Notes 5:107–109

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1122

Guizotia abyssinica (Niger)

Guizotia abyssinica (Niger) Family: Asteraceae

Oilseed

Cucumber mosaic virus

(CMV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV infection in plants of Guizotia abyssinica was reported from India (Sastry 1985; Jagadeeshwar and Madhusudan 1996). The virus-infected niger plants exhibit mosaic symptoms on the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Guizotia abyssinica was reported from Karnataka State, India (Arun Kumar et al. 2007). The virus-infected niger plants exhibit symptoms of leaf and petiole necrosis. The virus is transmitted by the thrips vectors (Frankliniella schultzei, Thrips palmi, and Thrips tabaci); the virus present in/on pollen entering the host through injuries caused by thrips while feeding. The virus is mechanically sap-transmissible and also by pollen. The virus is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

References Arun Kumar N, Lakshmi Narasu M, Zehr UB, Ravi KS (2007) First report of Tobacco streak virus infecting Guizotia abyssinica from India. Plant Dis 91(3):330 Jagadeeshwar R, Madhusudan T (1996) Identification and characterization of mosaic disease of niger. Plant Dis Res 11:178–179 Sastry KS (1985) A mosaic disease of niger (Guizotia abyssinica Cass.). J Oil-Seeds Res 3:144–145

Gymnaster savatieri (Miyakowasure) Family: Asteraceae

Ornamental

Chrysanthemum virus B Taxonomic position Genus: Carlavirus

(CVB)

Family: Betaflexiviridae

Gymnocalycium mihanovichii (Chin cactus)

1123

CVB infection in plants of Gymnaster savatieri was reported from Japan (Suastika et al. 1997). The virus-infected miyakowasure plants exhibit mild mottle symptoms on the leaves and color breaking of the flowers. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CVB, refer to Chrysanthemum spp.

References Suastika G, Kurihara J, Natsuaki KT, Tomaru K (1997) A strain of Chrysanthemum B carlavirus causing flower colour breaking on Gymnaster savatieri (Makino) Kitamura. Ann Phytopathol Soc Jpn 63:1–7

Gymnocalycium mihanovichii (Chin cactus) Family: Cactaceae

Ornamental

Cactus mild mottle virus Taxonomic position Genus: Tobamovirus

(CMMoV)

Family: Virgaviridae

Geographical distribution CMMoV infection in plants of Gymnocalycium mihanovichii was first reported from Korea (Min et al. 2006). Symptoms and host(s) The virus-infected chin cactus plants exhibit mild mottle symptoms in the form of rings along the stem. Transmission There is no known vector for this virus. The virus is mechanically sap-transmissible. The virus is grafttransmissible and also transmissible through contact between plants. Virion properties and genome The virions are non-enveloped, rigid helical rods with a helical symmetry, about 18 nm in diameter and 320 nm in length. The genome is a monopartite, linear, positive-sense ssRNA of 6449 nt (EU043335 = NC_011803). The 30 -terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G5’pppG) (Min et al. 2006, 2009; Zaitlin 2011; Adams et al. 2017).

References Adams MJ, Adkins S, Bragard C, Gilmer D, Li D, MacFarlane SA, Wong S-M, Melcher U, Ratti C, Ryu KH, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Virgaviridae. J Gen Virol 98:1999–2000 Min BE, Chung BN, Kim MJ, Ha JH, Lee BY, Ryu KH (2006) Cactus mild mottle virus is a new cactus-infecting tobamovirus. Arch Virol 151:13–21 Min BE, Song YS, Ryu KH (2009) Complete sequence and genome structure of cactus mild mottle virus. Arch Virol 154(8):1371–1374 Zaitlin M (2011) Tobamovirus. In: Tidona C, Darai G (eds) The Springer Index of Viruses. Springer, New York, NY, https://doi.org/10.1007/978-0-387-95919-1

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1124

Gynura aurantiaca (Purple passion, Velvet plant)

Gynura aurantiaca (Purple passion, Velvet plant) Family: Asteraceae

Citrus exocortis viroid Taxonomic position Genus: Pospiviroid

Ornamental

(CEVd)

Family: Pospiviroidae

CEVd infection in plants of Gynura aurantiaca was reported from Spain (Chaffai et al. 2007). The viroid-infected purple passion plants exhibit mild or severe symptoms on the foliage. The viroid is mechanically transmissible from plant to plant, and the use of planting material from infected plants is the primary source of viroid spread. For more details of CEVd, refer to Citrus spp.

References Chaffai M, Serra P, Gandía M, Hernández C, Duran-Vila N (2007) Molecular characterization of CEVd strains that induce different phenotypes in Gynura aurantiaca: structure-pathogenicity relationships. Arch Virol 152:1283–1294

Gynura procumbens (Longevity spinach) Family: Asteraceae

Broad bean wilt virus 2 Taxonomic position Genus: Fabavirus

Medicinal plant

(BBWV-2)

Family: Secoviridae

BBWV-2 infection in plants of Gynura procumbens was reported from Korea (Kwak et al. 2017). The virus-infected longevity spinach plants exhibit symptoms of chlorotic local lesions, vein chlorosis, mosaic and mottle on the leaves, and stunting of plants. The virus is transmitted by the aphid vector, Acyrthosiphon solani, in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. Virus spread takes place through the use of infected vegetative planting material. For more details of BBWV-2, refer to Vicia faba.

References Kwak H-R, Go W-R, Kim M, Kim C-S, Choi H-S, Seo J-K, Kim J-G, Kim J-S (2017) First report of Broad bean wilt virus 2 in Gynura procumbens in Korea. Plant Dis 101:514

Gypsophila spp. (Baby’s breath)

1125

Gypsophila spp. (Baby’s breath) Family: Caryophyllaceae

Ornamental

Carnation Italian ringspot virus Taxonomic position Genus: Tombusvirus

(CIRV)

Family: Tombusviridae

CIRV was reported in plants of Gypsophila paniculata from the Netherlands (Koenig et al. 2004, 2009). The virus-infected baby’s breath plants showed chlorotic and necrotic lesions. The virus spreads in water and soil without involvement of any vector. The virus is mechanically sap-transmissible and graft-transmissible, use of infected budwood helps spread this virus. For more details of CIRV, refer to Dianthus caryophyllus.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV was reported infecting Gypsophila paniculata with mosaic symptoms in Korea (Park and Choi 2009) and Japan. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Gypsophila paniculata with virus-like symptoms was reported from Lithuania, and ToRSV purified from such plants (Navalinskiene and Samuitiene 2000; Samuitiene and Navalinskiene 2001). The virus is transmitted by nematode vectors in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and also by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

G

1126

Gypsophila spp. (Baby’s breath)

TSWV infection in plants of Gypsophila elegans was reported from Zimbabwe (Karavina and Gubba 2017). The virus-infected baby’s breath plants exhibit symptoms of ringspots, line patterns, mottling and chlorotic blotches on leaves, severe stunting, wilting, and even plant death. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sapinoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Karavina C, Gubba A (2017) Detection and characterization of Tomato spotted wilt virus infecting field and greenhousegrown crops in Zimbabwe. Eur J Plant Pathol 149:933–944 Koenig R, Verhoeven JT, Fribourg CE, Pfeilstetter E, Lesemann D-E (2004) Evaluation of various species demarcation criteria in attempts to classify ten new tombusvirus isolates. Arch Virol 149:1733–1744 Koenig R, Lesemann DE, Pfeilstetter E (2009) New isolates of carnation Italian ringspot virus differ from the original one by having replication-associated proteins with a typical tombusvirus-like N-terminus and by inducing peroxisomerather than mitochondrion-derived multivesicular bodies. Arch Virol 154(10):1695–1698 Navalinskiene M, Samuitiene M (2000) Natural occurrence of tomato ringspot nepovirus in ornamental plants in Lithuania. Proceedings of the international conference: development of environmentally friendly plant protection in the Baltic Region, Tartu, Estonia, September 28–29. pp 140–143 Park H-S, Choi JK (2009) Characterization of Cucumber mosaic virus isolated from Gypsophila paniculata. J Agric Life Sci 21:51–58 Agriculture and Life Sciences Research Institute, Kangwon National University Samuitiene M, Navalinskiene M (2001) Nepoviruses and their influence in field horticulture. Biologija 4:43–45

H

Habenaria radiata (White egret flower) Synonyms Pecteilis radiata Family: Orchidaceae

Ornamental

Habenaria mosaic virus Taxonomic position Genus: Potyvirus

(HaMV)

Family: Potyviridae

Geographical distribution HaMV was first reported in plants of Habenaria radiata from Japan by Inouye (1983). The virus spreads in Japan (Inouye et al. 1998; Kondo et al. 2014).

Symptoms and host(s) The virus-infected white egret flower plants exhibit mosaic and conspicuous leaf chlorosis symptoms. Transmission The virus is transmitted by an aphid vector Myzus persicae in a non-persistent manner. The virus is transmissible by mechanical inoculation to several (3–9) families, causing only localized infection.

Virion properties and genome The virions are non-enveloped, flexuous filaments with a clear modal length of 750 nm and13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9499 nt (AB818538 = NC_021786) (Kondo et al. 2014; Revers and Garcia 2015; Wylie et al. 2017). © Springer Nature India Private Limited 2019 K. S. Sastry et al., Encyclopedia of Plant Viruses and Viroids, https://doi.org/10.1007/978-81-322-3912-3

1128

Halesia carolina (Carolina silverbell)

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV infection in plants of Habenaria radiata was reported from Japan (Gara et al. 1997). The virus-infected white egret flower plants exhibited symptoms of mosaic, leaf distortion, and severe stunting. The virus is transmitted by an aphid vector (Myzus persicae) in a non-persistent manner. The virus is readily transmissible by mechanical inoculation of sap to 20 of 52 plant species in 8 of 11 families. For more details of WMV, refer to Citrullus lanatus.

References Gara IW, Kondo H, Maeda T, Inouye N, Tamada T (1997) Stunt disease of Habenaria radiata caused by a strain of Watermelon mosaic virus 2. Ann Phytopathol Soc Jpn 63:113–117 Inouye N (1983) Habenaria mosaic virus, a new member of the potyvirus group. 4th Int. Cong. Plant Pathol, p. 117 Inouye N, Maeda T, Mitsuhata K, Gara IW (1998) Habenaria mosaic potyvirus, a new member of potyvirus from Habenaria radiata orchid. Bull Res Inst Bioresour Okayama Univ 5:155–168 Kondo H, Maeda T, Gara IW, Chiba S, Maruyama K, Tamada T, Suzuki N (2014) Complete genome sequence of Habenaria mosaic virus, a new potyvirus infecting a terrestrial orchid (Habenaria radiata) in Japan. Arch Virol 159:163–166 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354

Halesia carolina (Carolina silverbell) Family: Styracaceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Halesia carolina was reported from Georgia, USA (Ruter and Gitaitis 1993). The virus-infected Carolina silverbell plants were asymptomatic. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

References Ruter JM, Gitaitis RD (1993) Impatiens necrotic spot virus in woody landscape plants in Georgia. Plant Dis 77:318

Hardenbergia spp. (Hardenbergia violacea; H. comptoniana) (Vine lilac)

1129

Hardenbergia spp. (Hardenbergia violacea; H. comptoniana) (Vine lilac) Family: Fabaceae

Ornamental

Hardenbergia mosaic virus Taxonomic position Genus: Potyvirus

(HarMV)

Family: Potyviridae

Geographical distribution HarMV infection in plants of Hardenbergia spp. was reported from Australia (Webster et al. 2007; Luo et al. 2011; Wylie and Jones 2011a). Symptoms and host(s) The virus-infected vine lilac plants exhibit symptoms varying from mild mosaic to severe mosaic with leaf deformation. Transmission The virus is transmitted by aphid vectors, Myzus persicae, Acyrthosiphon kondoi, and Rhopalosiphum padi, in a non-persistent manner (Luo et al. 2011). The virus is transmissible by mechanical sap-inoculation to several species in the Amaranthaceae, Chenopodiaceae, Fabaceae, and Solanaceae. No seed transmission is observed. Virion properties and genome The virions are non-enveloped, flexuous filaments 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9682 nt (HQ161081 = NC_015394) (Kehoe et al. 2014; Wylie and Jones 2011a; Wylie et al. 2017).

Hardenbergia virus A Taxonomic position Genus: Divavirus

(HarVA)

Family: Betaflexiviridae

Geographical distribution HarVA infection in plants of Hardenbergia spp. was reported from Australia (Wylie and Jones 2011b). Symptoms and host(s) The virus-infected vine lilac plants exhibit mild mosaic symptoms Transmission No information on transmission is available. Virion properties and genome Particles are flexuous rods, encapsidating a single-stranded RNA genome of 6942 nt (NC_015395; HQ241409) (Wylie and Jones 2011b.

H

1130

Hebe spp.

References Kehoe MA, Coutts BA, Buirchell BJ, Jones RA (2014) Hardenbergia mosaic virus: crossing the barrier between native and introduced plant species. Virus Res 184:87–92 Luo H, Wylie SJ, Coutts B, Jones RAC, Jones MGK (2011) A virus of an isolated indigenous flora spreads naturally to an introduced crop species. Ann Appl Biol 159:339–347 Webster CG, Coutts BA, Jones RAC, Jones MGK, Wylie SJ (2007) Virus impact at the interface of an ancient ecosystem and a recent agroecosystem: studies on three legume infecting potyviruses in the southwest Australian floristic region. Plant Pathol 56:729–742 Wylie S, Jones M (2011a) Characterisation and quantitation of mutant and wild-type genomes of Hardenbergia mosaic virus isolates co-infecting a wild plant of Hardenbergia comptoniana. Arch Virol 156(7):1251–1255 Wylie S, Jones M (2011b) Hardenbergia virus A, a novel member of the family Betaflexiviridae from a wild legume in Southwest Australia. Arch Virol 156(7):1245–1250 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354

Hebe spp. Family: Plantaginaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Ornamental

(AMV)

Family: Bromoviridae

AMV was isolated from plants of Hebe spp. but no symptoms were reported (Schwenk et al. 1969). The virus is transmitted by a large number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

References Schwenk FW, Williams HE, Smith HS (1969) Alfalfa mosaic virus from Hebe, Ilex and Viburnum. Phytopathology 59:1048–1049

Hedera helix (Ivy) Family: Araliaceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

Family: Secoviridae

Hedera helix (Ivy)

1131

ArMV was consistently associated with Hedera helix plants showing chlorotic yellow vein-banding, blotches, or broad ring patterns in the United Kingdom (Cooper and Sweet 1976). The virus is transmitted by nematode vectors (Xiphinema spp.) in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Ivy vein banding virus

(IVBV)

Taxonomic position IVBV is a tentative member of the genus Cytorhabdovirus and family Rhabdoviridae Geographical distribution IVBV infection in plants of Hedera helix was reported from the Czech Republic (Petrzik 2012). Symptoms and host(s) The virus-infected ivy plants exhibit symptoms ranged from mild mosaic to vein-clearing, vein yellowing, and leaf curling. Virion properties and genome The virions are bacilliform or bullet-shaped particles of 60–70  150–300 nm; partial genomic sequences are available (e.g., GQ249162, GQ249163) (Walker et al. 2018).

Ivy vein clearing virus

(IVCV)

Taxonomic position IVCV is a tentative member of the family Rhabdoviridae Geographical distribution IVCV was first reported in plants of Hedera helix from southern Italy by Russo et al. (1979). The virus spreads in Italy and Yugoslavia (Albouy and Devergne 1999). Symptoms and host(s) The virus-infected ivy plants exhibit transitory chlorosis of leaf veins, with some chlorotic blotching in spring. Transmission The virus is transmissible by mechanical inoculation to several plant families (Castellano and Rana 1981). Virion properties and genome The virions are 55 nm in diameter and 325 nm long.

References Albouy J, Devergne JC (1999) Diseases produced by viruses on ornamental plants. Ediciones Mundi-Prensa, Madrid 480 p Castellano MA, Rana GL (1981) Transmission and ultrastructure of Ivy vein clearing virus infections. Phytopathol Mediker 20:199–202

H

1132

Hedyotis uncinella

Cooper JI, Sweet JB (1976) The detection of viruses with nematode vectors in six woody hosts. Forestry 49:73–78 Petrzik K (2012) Bioinformatic analysis of the L polymerase gene leads to discrimination of new rhabdoviruses. J Phytopathol 160:377–381 Russo M, Castellano MA, Martelli GP (1979) Rhabdovirus-like particles in English ivy (Hedera helix L.) and ivy-leafed geranium (Pelargonium peltatum). Phytopath Z 96:122–131 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, and ICTV Report Consortium (2018) ICTV Virus Taxonomy Profile: Rhabdoviridae. J Gen Virol 99:447–448

Hedyotis uncinella Family: Rubiaceae

Ornamental

Hedyotis uncinella yellow mosaic virus Taxonomic position Genus: Begomovirus

(HeuYMV)

Family: Geminiviridae

Geographical distribution HeuYMV infection in plants of Hedyotis uncinella was reported from Vietnam (Du et al. 2014). Symptoms and host(s) The virus-infected hedyotis plants exhibit yellow mosaic symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2749 nt (KF429251) (Briddon 2001; Du et al. 2014; Brown et al. 2015; Zerbini et al. 2017). Hedyotis uncinella yellow mosaic betasatellite DNA molecule is associated with HeuYMV, and consists of 1348 nt (KF641186) (Zhou 2013; Du et al. 2014).

References Briddon RW (2001) Begomovirus. Geminiviridae. The Springer Index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Du Z, Chen M, Wang Z, Liu Y, Zhang S, He Z (2014) Isolation and molecular characterization of a distinct begomovirus and its associated betasatellite infecting Hedyotis uncinella (Hook. et Arn.) in Vietnam. Virus Genes 48(3):557–561

Helenium spp. (Sneezeweed)

1133

Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zhou X (2013) Advances in understanding begomovirus satellites. Annu Rev Phytopathol 51:357–381

Helenium spp. (Sneezeweed) Family: Asteraceae

Helenium virus S Taxonomic position Genus: Carlavirus

Ornamental

(HVS)

Family: Betaflexiviridae

Geographical distribution HVS was first reported in plants of Helenium amarum from Germany by Kuschki et al. (1978). The virus spreads in Germany and the USA (Foster et al. 1990). Symptoms and host(s) The virus-infected sneezeweed plants exhibit symptomless infections. Infection has also been reported from Impatiens spp. and Veronica spp. Transmission The virus is transmitted by the aphid vector Myzus persicae in a non-persistent manner. The virus is transmissible by mechanical inoculation to less than three plant families. The virus is not transmitted by seed. Virion properties and genome The virions are flexuous filaments about 610–700 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA 7.4–7.9 kb in size and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Foster et al. 1990; Adams et al. 2004).

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Foster GD, Millar AW, Meehan BM, Mills PR (1990) Nucleotide sequence of the 30 -terminal region of Helenium virus S RNA. J Gen Virol 71(PT 8):1877–1880 Kuschki GH, Koenig R, Düvel D, Kühne H (1978) Helenium virus S and Y – two new viruses from commercially grown Helenium hybrids. Phytopathology 68:1407–1411

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1134

Helianthus annuus (Sunflower)

Helianthus annuus (Sunflower) Family: Asteraceae

Oilseed

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

ArMV infection in plants of Helianthus annuus was reported from Iran (Ghotbi and Shahraeen 2005). The virus-infected sunflower plants exhibit mosaic, leaf chlorosis, small necrotic lesions, and leaf malformation and deformation symptoms. This virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Beet curly top virus Taxonomic position Genus: Curtovirus

(BCTV)

Family: Geminiviridae

BCTV infection in plants of Helianthus annuus was reported from California, USA (Creamer et al. 1996). The virus is transmitted by a leafhopper vector, Circulifer tenellus, in a persistent (circulative, non-propagative) manner. The virus is not transmissible by mechanical inoculation. For more details of BCTV, refer to Beta vulgaris.

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

BWYV infection in plants of Helianthus annuus was reported from the UK (Russel et al. 1975; Tomlinson 1988). The virus-infected sunflower plants exhibited pale yellow zones appearing between the veins on infected leaves, followed by necrotic spots on leaf tips and margins. The virus is aphid transmitted by Aphis craccivora, A. gossypii, Myzus persicae, and other aphid species, in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation, and is also not transmitted by seed. For more details of BWYV, refer to Beta vulgaris.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

Helianthus annuus (Sunflower)

1135

CMV infection in plants of Helianthus annuus is cosmopolitan in distribution (Venugopal Rao et al. 1987; Mohan 1992; Nishimura et al. 1995; Rabiee et al. 2015). CMV infects sunflower in nature causing mosaic symptoms (Smith 1972). Orellana and Quacquarelli (1968) described a sunflower mosaic caused by the CMV-SF strain with symptom of mosaic and chlorotic rings on young leaves, increasing in severity on older leaves. Pale brown spots occur along the stalks and leaf petioles, growth of plants is stunted. The virus is transmitted by aphid vectors Aphis gossypii and Myzus persicae in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Groundnut bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(GBNV)

Family: Tospoviridae

GBNV infection in plants of Helianthus annuus was reported from India (Pranav et al. 2008; Jain et al. 2000). The virus-infected sunflower plants exhibit symptoms of necrosis of leaf and petiole, stunting of plant, malformation of the heads, and poor seed set. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of GBNV, refer to Arachis hypogaea.

Pelargonium zonate spot virus Taxonomic position Genus: Anulavirus

(PZSV)

Family: Bromoviridae

PZSV infection in plants of Helianthus annuus was reported from Argentina (Giolitti et al. 2013). The virus-infected sunflower plants exhibit chlorotic concentric rings and line patterns on the leaves. Thrips are involved in transmitting this virus by carrying the virus-infected pollen grains on their bodies and introduce the virus into wounds on the susceptible hosts while feeding. The virus is mechanically saptransmissible to a wide range of test plants. For more details of PZSV, refer to Pelargonium spp.

Potato virus Y

(PVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

PVY infection in plants of Helianthus annuus was reported from the Czech Republic and Iran (Chod et al. 1990, 1996; Rabiee et al. 2015). The virus-infected sunflower plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts and by grafting. For more details of PVY, refer to Solanum tuberosum.

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1136

Helianthus annuus (Sunflower)

Sunflower chlorotic mottle virus Taxonomic position Genus: Potyvirus

(SCMoV)

Family: Potyviridae

Geographical distribution SCMoV infection in plants of Helianthus annuus was reported from Argentina (Lenardon 1994; Dujovny et al. 1998; Formento et al. 1994; Lenardon et al. 2001; Giolitti et al. 2010). Two strains have been isolated from the main sunflower crop region: C (common) strain and RS (ringspot) strain. Symptoms and host(s) The virus-infected sunflower plants exhibit characteristic chlorotic ringspots, mosaic, and mottling symptoms. Mottling symptoms coalesce to yellow blotches followed by distortion and stunting of the plant (Schneiter and Miller 1981; Lenardon 1994). Transmission The virus is transmitted by aphid species in a non-persistent manner; Myzus persicae is the most efficient vector for this virus (Dujovny et al. 2000). The virus is also mechanically sap-transmissible, and host range included members of the Amaranthaceae, Asteraceae, Chenopodiaceae, and Solanaceae families. The virus is not seed-transmitted. Virion properties and genome The virions are non-enveloped, flexuous filaments 770 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9965 nt (GU181199 = NC_014038) (Dujovny et al. 1998; Bejerman et al. 2010; Revers and Garcia 2015; Wylie et al. 2017).

Sunflower crinkle virus

(SuCV)

Taxonomic position SuCV is a tentative member of the genus Umbravirus and family Tombusviridae Geographical distribution The virus was first reported in plants of Helianthus annuus from Kenya by Martens et al. (1970) and is also reported from Tanzania (Theuri et al. 1987). Symptoms and host(s) On sunflower (SuCV) produced blotch symptoms on turned into leaf crinkling symptoms. Transmission The virus is transmitted by the aphid vector Aphis gossypii (Theuri et al. 1987). The virus is mechanically transmissible to many plant species belonging to families Compositae (Asteraceae), Solanaceae, and Leguminosae. No seed transmission is reported. Virion properties and genome The virions measure 26 nm in diameter (Theuri et al. 1987).

Helianthus annuus (Sunflower)

1137

Sunflower mild mosaic virus Taxonomic position Genus: Potyvirus

(SMMV)

Family: Potyviridae

Geographical distribution SMMV infection in plants of Helianthus annuus was reported from Argentina (Lenardon 1994; Giolitti et al. 2015). Symptoms and host(s) The virus-infected sunflower plants exhibit chlorotic mild mosaic symptoms. Transmission The virus is transmitted by an aphid vector Myzus persicae in a non-persistent manner. The virus is also mechanically sap-transmissible. The virus is not transmitted by seed or dodder (Lenardon 1994). Virion properties and genome The virions are non-enveloped, flexuous filaments 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9718 nt (JQ350738 = NC_021065) (Giolitti et al. 2015; Revers and Garcia 2015; Wylie et al. 2017).

Sunflower mosaic virus Taxonomic position Genus: Potyvirus

(SuMV)

Family: Potyviridae

Geographical distribution SuMV infection in wild plants of Helianthus annuus was first reported from Texas, USA (Arnott and Smith 1967). The virus occurs wherever sunflower is grown (Gupta 1981; Nagaraju et al. 1997; Gulya et al. 1998, 2002). Symptoms and host(s) The virus causes mild to severe systemic symptoms in sunflower. There are also reports that SuMV infection reduced the plant height, stem girth, leaf area, head size, and seed weight of sunflower hybrids (Jindal et al. 2001; Gulya et al. 2002; Verma et al. 2009). Transmission The virus is transmitted by aphid vectors Myzus persicae and Capitphorus elaegni in a non-persistent manner. Seed transmission is reported to be up to 12% in sunflower. Virion properties and genome The virions are non-enveloped, flexuous filaments 720 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA. A partial genome sequence of 2646 nt is available (AF465545) (Gulya et al. 2002; Revers and Garcia 2015; Wylie et al. 2017).

H

1138

Helianthus annuus (Sunflower)

Sunflower ring blotch virus Taxonomic position Genus: Potyvirus

(SuRBV)

Family: Potyviridae

Geographical distribution SuRBV infection in plants of Helianthus annuus was reported from Argentina (Mederos et al. 2017). Symptoms and host(s) The virus-infected sunflower plants exhibit ring blotch symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. Virion properties and genome Virions are non-enveloped, flexuous filaments, 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 9555 nt in length (KX856009 = NC_034208) (Mederos et al. 2017; Wylie et al. 2017).

Sunflower yellow blotch virus

(SuYBV)

Taxonomic position SuYBV is a tentative member of the genus Umbravirus and family Tombusviridae Geographical distribution SuYBV infection in plants of Helianthus annuus was reported from Kenya, Zambia, and Tanzania (Theuri et al. 1987; Theuri and Njuho 1989; Ndunguru and Mwale 1999). Symptoms and host(s) The virus-infected sunflower plants exhibit blotching and leaf curling symptoms. Transmission The virus is transmitted by an aphid vector, Aphis gossypii. The virus is mechanically sap-transmissible to many different plant species of Compositae (Asteraceae), Solanaceae, and Leguminosae. The virus is not seed-transmitted. Virion properties and genome The virions measure 28 nm in diameter. The genome is a linear, positive-sense, single-stranded RNA.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

Helianthus annuus (Sunflower)

1139

TMV infection in plants of Helianthus annuus was reported from Egypt and New Zealand (Fletcher 1989; Zein et al. 2012). The virus-infected sunflower plants exhibit symptoms of yellow blotching of leaves and severe stunting of plants. There is no known vector for this virus. The virus is mechanically sap-transmissible and also through contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco rattle virus

(TRV)

Taxonomic position Genus: Tobravirus

Family: Virgaviridae

TRV infection in plants of Helianthus annuus was reported from Central Europe (Schmelzer 1970). The virus-infected sunflower plants show discoloration and mild necrosis of infected leaves. The virus is transmitted by nematode vectors Trichodorus spp. and is also transmissible by mechanical sap-inoculation to a number of hosts. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Helianthus annuus was reported from Texas, USA (McLean 1962). The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

Geographical distribution TSV infection in plants of Helianthus annuus is distributed worldwide, including the Pacific region, India, Australia, Canada, Peru, and the USA (Dijkstra 1983; Prasada Rao et al. 2000; Ramaiah et al. 2001; Ravi et al. 2001; Bhat et al. 2002a, b; Jain et al. 2003; Suresh et al. 2004; Lavanya et al. 2005; Lokesh et al. 2008; Sharman et al. 2008, 2015; Pankaja et al. 2010, 2011; Swamy et al. 2010; Hosseini et al. 2012; Papaiah et al. 2012). Symptoms and host(s) TSV-infected sunflower plants exhibit necrosis of the leaf lamina followed by twisting of the leaf and systemic mosaic/necrosis of the lamina, petiole, stem and floral calyx, and corolla, eventually resulting in dieback. Early infected plants remain stunted with malformed heads having chaffy or deformed seeds. Heads are partially twisted and become sterile (Pankaja et al. 2013).

H

1140

Helianthus annuus (Sunflower)

TSV affects a large number of crops, including bean, sunflower, peanut, maize, and soybean, and a range of vegetable and ornamental species. The weed Parthenium hysterophorus is a potential reservoir of infection. Transmission The virus is transmitted by thrips vectors, Frankliniella schultzei, Scirtothrips dorsalis, Megalurothrips usitatus, and Microcephalothrips abdominalis, which colonize the flowers of sunflower, and have been shown to serve as vectors and carry the infected pollen to the host plants. Thrips, while feeding, not only cause injury to leaf tissue but also deposit pollen and facilitate virus transmission to the plant (Sdoodee and Teakle 1987; Prasada Rao et al. 2003; Singh 2005). The virus is transmissible by mechanical sapinoculation, and by grafting but is not transmissible by contact between plants. No seed transmission of TSV was detected in a number of sunflower cultivars tested (Reddy et al. 2007; Bhat et al. 2013). However the virus is transmitted by pollen to the pollinated plant (Ajith-Prasad and Nagaraju 2005). Virion properties and genome The virions are isometric; non-enveloped; 27, 30, and 35 nm in diameter; rounded in profile; and without a conspicuous capsomere arrangement (Ramaiah et al. 2001). The virus has a tripartite genome, RNA1 (NC_003844), RNA2 (NC_003845), and RNA3 (NC_003845), acting as mRNAs. RNA3 carries the ORF, which codes for the viral coat protein, which is expressed from a subgenomic RNA. Sequence comparison of RNA3 and the coat protein gene provided evidence for the taxonomic position of TSV from sunflower (Ravi et al. 2001; Bhat et al. 2002b, c; Scott 2011a, b).

Tomato leaf curl Karnataka virus Taxonomic position Genus: Begomovirus

(ToLCKaV)

Family: Geminiviridae

ToLCKaV infection in plants of Helianthus annuus was reported from Karnataka (India) (Vanitha et al. 2013a, b; Vindyashree et al. 2014). The virus-infected sunflower plants exhibit symptoms of upward curling and reduction in leaf size with prominent enations on the lower surface of the leaves followed by severe stunting and bushy appearance with poor earhead formation. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. The virus was successfully transmitted from sunflower to H. annuus, Datura stramonium, and Parthenium hysterophorus by means of Bemisia tabaci. The virus is graft-transmissible. For more details of ToLCKaV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Helianthus annuus is cosmopolitan in distribution (Granoff and Webster 1999; Rabiee et al. 2015). The virus-infected sunflower plants exhibit yellow-to-pale green interveinal mottle occurring on the dwarfed, malformed, infected leaves; ringspots occur on stalks. Grooves and fissures form on the upper parts of the stalks and on leaf petioles, and the vascular tissues turn black. Internodes shorten and flower heads do not form or remain small and bent over. The virus is transmitted

Helianthus annuus (Sunflower)

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by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Ajith-Prasad HN, Nagaraju (2005) Transmission of Sunflower necrosis virus disease through sap, seed, insect vector and pollen. Environ Ecol 23:125–128 Arnott HJ, Smith KM (1967) Electron microscopy of virus infected sunflower leaves. J Ultrastruct Res 25:3389–3402 Bejerman N, Giolitti F, de Breuil S, Lenardon S (2010) Molecular characterization of Sunflower chlorotic mottle virus: a member of a distinct species in the genus Potyvirus. Arch Virol 155(8):1331–1335 Bhat BN, Raja Ram Reddy D (2013) Studies on seed transmission of Tobacco streak virus causing sunflower necrosis disease. Int J Plant Prot 6(1):48–50 Bhat AI, Jain RK, Chaudhury V, Krishnareddy M, Ramaiah K, Chattannavar SN, Varma A (2002a) Sequence conservation in the coat protein gene of Tobacco streak virus isolates causing necrosis disease in cotton, mungbean, sunflower, and sunn-hemp in India. Indian J Biotechnol 1:350–356 Bhat AI, Jain RK, Kumar A, Ramiah M, Varma A (2002b) Serological and coat protein sequences studies suggest that necrosis disease of sunflower in India is caused by a strain of Tobacco streak ilarvirus. Arch Virol 147:651–658 Bhat AI, Jain RK, Ramiah M (2002c) Detection of Tobacco streak virus from sunflower and other crops by reverse transcription polymerase chain reaction. Indian Phytopathol 55:216–218 Chod J, Skaloud V, Jokes M (1990) Detection of potato Y virus in connection with sunflower mosaic symptoms. Sb UVTIZ Ochrana Rostlin 26:11–16 Chod J, Skaloud V, Jokes M (1996) Detection of Potato Y virus in the connection with sunflower mosaic symptoms. Sb. UVITZ Ochrana Rostlin 26:11–16 Creamer R, Lugue-Williams M, Howo M (1996) Epidemiology and incidence of Beet curly top geminivirus in naturally infected weed hosts. Plant Dis 80:533–535 Dijkstra J (1983) Tobacco streak virus in sunflower (Helianthus annuus). Neth J Plant Pathol 89:153–169 Dujovny G, Usugi T, Shohara K, Lenardon SL (1998) Characterization of a potyvirus infecting sunflower in Argentina. Plant Dis 82:470–474 Dujovny G, Sasaya T, Koganezawa H, Usugi T, Shohara K, Lenardon SL (2000) Molecular characterization of a new Potyvirus infecting sunflower. Arch Virol 145:2249–2258 Fletcher JD (1989) Additional hosts of Alfalfa mosaic virus, Cucumber mosaic virus and Tobacco mosaic virus in New Zealand. N Z J Crop Hortic Sci 17:361–362 Formento N, Rodriguez Pardina P, Alegre A, Truol G, Laguna G (1994) Potyvirus, causal organism of sunflower mosaic in Argentina. Fitopatologia 29:36–37 Ghotbi T, Shahraeen N (2005) First report on incidence of Arabis mosaic virus (ArMV, Nepovirus) on ornamental plants in Iran. Iran J Plant Pathol 41(2):305–306 Giolitti FJ, Bejerman NE, De Breuil S, Lenardon SL (2010) Identification and characterization of a new strain of sunflower chlorotic mottle virus, a potyvirus infecting Asteraceae in Argentina. J Phytopathol 158:536–541 Giolitti F, Bejerman N, Nome C, Visintin G, de Breuil S, Lenardon S (2013) Biological and molecular characterization of an isolate of Pelargonium zonate spot virus infecting sunflower in Argentina. J Plant Path 96:189–194 Giolitti F, de Breuil S, Bejerman N, Lenardon S (2015) Complete nucleotide sequence and genetic organization of Sunflower mild mosaic virus (SuMMoV). Sourced from asagir.org.ar, April 14 2015 Granoff A, Webster RG (1999) Encyclopedia of virology, vol I–III. Academic, New York Gulya TJ, Berger PH, Shiel PJ, Freeman TF, Isakeit TA, Berger PH (1998) Mosaic – causing potyvirus on wild sunflower from southern Texas. Phytopathology 88:S34 Gulya TJ, Shiel PJ, Freeman T, Jordan RL, Isakeit T, Berger PH (2002) Host range and characterization of Sunflower mosaic virus. Phytopathology 92:694–702 Gupta KC (1981) Studies on the identity of Sunflower mosaic virus. In: 3rd Int. Symp. on Plant Pathol New Delhi, December 14–18. p 117 Hosseini S, Koohi Habibi M, Mosahebi G, Motamedi M, Winter S (2012) First report on the occurrence of Tobacco streak virus in sunflower in Iran. J Plant Pathol 94:585–589 Jain RK, Bhat AI, Byadgi AS, Nagaraju J, Singh H, Halkeri AV, Anahosur KH, Varma A (2000) Association of a tospovirus with sunflower necrosis disease. Curr Sci 79:1703–1705 Jain RK, Bhat AI, Varma A (2003) Sunflower necrosis disease-an emerging viral problem, Technical Bulletin-I. Unit of Virology, IARI, New Delhi 11 pp

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Helianthus annuus (Sunflower)

Jindal S, Cheema SS, Kang SS (2001) Effect of Sunflower mosaic virus on vigour and productivity in sunflower hybrids. Plant Dis Res 16:79–83 Lavanya NM, Ramiah R, Sankaralingam A, Renukadevi P, Velazhahan R (2005) Identification of hosts for ilarvirus associated with sunflower necrosis disease. Acta Phytopathol Entomol Hung 40:31–34 Lenardon SL (1994) Sintomas de etiologia viral en cultivos de girasol. En Enfermedades del Girasol en la Argentina. 1994. Capitulo 15. Editores Pereyra VR y Escande AR. 113 pages Lenardon SL, Giolitti F, Leon AJ, Bazzalo ME, Grondona M (2001) Effects of Sunflower chlorotic mottle virus infections on sunflower yield parameters. Helia 24:55–66 Lokesh BK, Maraddi GN, Agnal MG, Upperi SN (2008) Present status of different modes of transmission of Sunflower necrosis virus (SNV) on sunflower weeds and crop plants. Int J Plant Protect 1(2):25–28 Martens JW, Ravagan M, McDoland WC (1970) Diseases of sunflower in Kenya. East Afr Agric For J 35:103–106 McLean DM (1962) Common weeds of Tobacco ring spot virus in the lower Rio Grande Valley of Texas. Plant Dis Reptr 48:5–7 Mederos DC, Bejerman N, Trucco V, de Breuil S, Lenardon S, Giolitti F (2017) Complete genome sequence of Sunflower ring blotch virus, a new Potyvirus infecting sunflower in Argentina. Arch Virol 162(6):1787–1790 Mohan J (1992) Identification and epidemiology of viruses affecting oilseeds crop in western UP. With special reference to mustard and sunflower mosaic, Final progress report of the project from May, 7, 1988 to May 6, 1991. Janata Vedic College, Meerut University, Baraut, p 30 Nagaraju V, Muniyappa V, Singh SJ, Virupakshappa K (1997) Occurrence of mosaic virus disease on sunflower in Karnataka. Indian Phytopathol 50:277–281 Ndunguru J, Mwale S (1999) Putative virus disease of sunflower in Zambia. EPPO Bull 29:215–216 Nishimura NT, Kikuchi S, Tsuda A, Tsuchizaki T (1995) Cucumber mosaic virus isolated from sunflower (Helianthus annuus L.). Proc Kanto Tosan Plant Prot Soc 42:151–152 Orellana RG, Quacquarelli AA (1968) Sunflower mosaic caused by a strain of Cucumber mosaic virus. Phytopathology 58:1439–1440 Pankaja NS, Harish Babu GV, Nagaraju (2010) Virus vector relationship studies of Sunflower necrosis virus (SNV) and its vector Thrips palmi (Karny). Int J Plant Protect 3(2):260–263 Pankaja NS, Harish Babu GV, Nagaraju (2011) Transmission modes of Sunflower necrosis virus determination and confirmation. Int J Plant Protect 4(1):38–42 Pankaja NS, Nagaraju, Harish Babu GV (2013) Studies on host range of Sunflower necrosis virus. J Plant Dis Sci 8:160–165 Papaiah S, Sai Gopal DVR, Sastry KS, Narasimha G (2012) Symptomological and biochemical studies on sunflower necrosis disease in sunflower plants in Rayalaseema region of Andhra Pradesh, India. Ann Biol Res 3:170–178 Pranav C, Krishnaraj PU, Kuruvinashetty MS (2008) Identification of Peanut bud necrosis virus in sunflower. J Plant Dis Sci 3:56–59 Prasada Rao RDVJ, Reddy AS, Chander Rao S, Varaprasad KS, Thirumala Devi K, Nagaraju, Muniyappa V, Reddy DVR (2000) Tobacco Streak ilarvirus as causal agent of sunflower necrosis disease in India. J Oilseeds Res 17:400–401 Prasada Rao RDVJ, Reddy AS, Reddy SV, Thirumala Devi K, Chander Rao S, Manoj Kumar V, Subramaniam K, Yellamanda Reddy T, Nigam SN, Reddy DVR (2003) The host range of Tobacco streak virus in India and transmission by thrips. Ann Appl Biol 142:365–368 Rabiee S, Hosseini S, Hosseini A (2015) Occurrence and distribution of some sunflower viruses from sunflower fields in Kerman and Isfahan provinces, Iran. Arch Phytopathol Plant Protect 48:223–228 Ramaiah M, Bhatt AI, Jain RK, Pant RP, Ahlawat YS, Prabhakar K, Varma A (2001) Isolation of an isometric virus causing sunflower necrosis in India. Plant Dis 85:443.2 Ravi KS, Butt Gereitt A, Kitkaru AS, Deshmukh S, Lesemann DE, Winter S (2001) Sunflower necrosis disease from India is caused by an ilarvirus related to tobacco streak virus. Plant Pathol 50:800 Reddy AS, Subramanyam K, Kumar PL, Waliyar F (2007) Assessment of Tobacco streak virus (TSV) transmission through seed in groundnut and sunflower. J Mycol Plant Pathol 37:136–137 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Russell GE, Cook PHL, Bunting ES (1975) An aphid-transmitted yellowing disease of sunflower. Plant Pathol 24:58–59 Schmelzer K (1970) Die Ursache des krauselmosaiks an Sonnenblumen. Nachrichtenbl Dt Pflanzens Chutzd (Berlin) NF 24:1–5 Schneiter AA, Miller JF (1981) Description of sunflower growth stages. Crop Sci 21:901–903 Scott SW (2011a) Bromoviridae and allies. In: Encyclopedia of life sciences (ELS). Wiley, Chichester. https://doi.org/ 10.1002/9780470015902 Scott SW (2011b) Ilarvirus. Bromoviridae. In: The Springer Index of Viruses. Springer, New York, pp 187–194. https:// doi.org/10.1007/978-0-387-95919-1_27 Sdoodee R, Teakle DS (1987) Transmission of Tobacco streak virus by Thrips tabaci: a new method of plant virus transmission. Plant Pathol 36:377–380 Sharman M, Thomos JE, Persley DM (2008) First report of Tobacco streak virus in sunflower (Helianthus annuus), cotton (Gossypium hirsutum), chickpea (Cicer arietinum), mungbean (Vigna radiata) in Australia. Aust Plant Dis Notes 3:27–29

Helichrysum spp.

1143

Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct Tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207 Singh H (2005) Thrips incidence and necrosis disease in sunflower (Helianthus annuus L.). J Oilseeds Res 22:90–92 Smith KM (1972) A text book of plant virus diseases, 3rd edn. Academic, New York 684 pp Suresh K, Kulkarni VR, Byadgi AS (2004) Detection and survey for the incidence of Tobacco streak virus causing sunflower necrosis disease in northern Karnataka. Indian Phytopathol 57:337 Swamy L, Katti P, Patil BV, Prabhuraj A, Chandranath HT (2010) Transmission and serological detection of Sunflower necrosis virus through insect vector, Thrips palmi Karny. Karnataka J Agric Sci 23(1):118–120 Theuri JM, Njuho PM (1989) A note on the effect of yellow blotch virus on yield of sunflower in Kenya. E Afr Agric For J 55:41–43 Theuri JM, Bock KR, Woods RD (1987) Distribution, host range and some properties of a virus disease of sunflower in Kenya. Trop Pest Manag 33:202–207 Tomlinson JA (1988) Beet western yellows virus (BWYV). In: Smith IM et al (eds) European handb Pl Dis. Blackwell Sci Publ, London, pp 21–23 Vanitha LS, Shankarappa KS, Rangaswamy KT, Wickramaarachchi WART, Govindappa MR (2013a) Complete nucleotide sequence of Tomato leaf curl Karnataka virus and ß satellite molecule associated with leaf curl disease on sunflower in India. Plant Pathol J 12:19–25 Vanitha LS, Rangaswamy KT, Govindappa MR, Manjunatha L, Shivakumar SC, Govardhana M (2013b) Survey, vector relationships, host range studies of Tomato leaf curl Karnataka virus causing sunflower leaf curl disease. Trends Biosci 6:36–39 Venugopal Rao R, Madhusudhan T, Sastry KS (1987) Studies on a mosaic disease of sunflower. J Oilseeds Res 4:286–288 Verma KP, Thakur MP, Dantre RK (2009) Occurrence of Sunflower mosaic virus and its effect on yield contributing characters of sunflower. J Interacademicia 13:245–246 Vindyashree M, Govindappa MR, Ghante VN, Aswathanarayana, Shankar Goud DS (2014) Host range and virus vector relationships of leaf curl begomovirus diseases on sunflower in relation to disease epidemiology. 23rd national conference on “Recent trends in virology Research in the Omics and Era” held at Coimbatore, Tamilnadu, p 149. Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Zein SN, El-Khalik A, Samaa K, Eman AAH, Azzam CR (2012) Characterization of Tobacco mosaic virus (TMV-S) isolated from sunflower (Helianthus annuus L.) in Egypt. Int J Virol 8:27–38

Helichrysum spp. Family: Asteraceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV was reported infecting plants of Helichrysum bracteatum in Portugal (Louro 1996). The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV was reported infecting plants of Helichrysum bracteatum from Bulgaria, Portugal, and the USA (Hausbeck et al. 1992; Louro 1996; Bakardjieva et al. 1998). The virus-infected helichrysum plants

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Heliconia spp.

exhibit necrosis and chlorotic spot symptoms. The virus is transmitted by thrips vectors in a persistentpropagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Helichrysum bracteatum was reported from Bangladesh (Ara et al. 2012). The virus-infected helichrysum plants exhibit mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

References Ara MR, Masud MMH, Akanda AM (2012) Detection of plant viruses in some ornamental plants that act as alternate hosts. Agriculturists 10:46–54 Bakardjieva N, Denkova S, Hristova D (1998) Tomato spotted wilt virus on ornamental species in Bulgaria. Biotech Biotechnol Equip 12:49–52 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among greenhouse ornamentals in Pennsylvania. Plant Dis 76:795–800 Louro D (1996) Detection and identification of tomato spotted wilt virus and Impatiens necrotic spot virus in Portugal. Acta Hortic 431:99–105

Heliconia spp. Family: Heliconiaceae

Ornamental

Banana bunchy top virus Taxonomic position Genus: Babuvirus

(BBTV)

Family: Nanoviridae

BBTV infection in plants of Heliconia spp. was reported from Hawaii (Hamim et al. 2017). The virus-infected heliconia plants exhibit symptoms of chlorotic leaves and green flecking of the veins. The virus is transmitted by the banana black aphid, Pentalonia nigronervosa, in a persistent but not in a propagative manner. The virus is not mechanically sap-transmissible. For more details of BBTV, refer to Musa spp.

References Hamim I, Green JC, Borth WB, Melzer MJ, Wang YN, Hu JS (2017) First report of Banana bunchy top virus in Heliconia spp. on Hawaii. Plant Dis 101:2153

Helleborus spp.

1145

Helleborus spp. Family: Ranunculaceae

Ornamental

Broad bean wilt virus 2 Taxonomic position Genus: Fabavirus

(BBWV-2)

Family: Secoviridae

BBWV-2 infection in plants of Helleborus spp. was reported from Germany and Scotland (Murant and Roberts 1977; Kleinhempel 1991a). The virus-infected helleborus plants exhibit ringspot symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BBWV-2, refer to Vicia faba.

H Carnation latent virus Taxonomic position Genus: Carlavirus

(CLV)

Family: Betaflexiviridae

CLV was reported in plants of Helleborus spp. in Germany (Richert-Poggeler et al. 2015). The virus is transmitted by the aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of CLV, refer to Dianthus caryophyllus.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV was reported in plants of Helleborus niger from Germany (Kleinhempel 1991b) and H. foetidus (bear’s foot) in France and Italy; yellow-to-white oak leaf and line patterns were observed in leaves of infected H. foetidus plants (Cardin et al. 2003). The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Helleborus mosaic virus Taxonomic position Genus: Carlavirus

(HeMV)

Family: Betaflexiviridae

Geographical distribution HeMV was first reported in to be widespread in plants of Helleborus niger from Germany by Koenig (1985).

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Helleborus spp.

Symptoms and host(s) The virus-infected helleborus plants exhibit symptoms of chlorotic and/or necrotic lesions along leaf veins. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are flexuous filaments about 610–700 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA 7.4–7.9 kb in size and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. A 30 partial sequence of 3030 nt is available (FJ196838). Coat protein subunits are of one type and 31–36 kDa in size (Adams et al. 2004).

Helleborus net necrosis virus Taxonomic position Genus: Carlavirus

(HNNV)

Family: Betaflexiviridae

Geographical distribution HNNV infection in plants of Helleborus spp. was reported from Europe, the USA, Japan, New Zealand, and Germany (Eastwell et al. 2009; Liefting et al. 2010; Gera et al. 2011; Shiraishi et al. 2011; RichertPoggeler et al. 2015). Symptoms and host(s) The virus-infected helleborus plants exhibit black spots or streaks on the leaves, stem, and flowers. The virus-infected plants become stunted and will die. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are flexuous filaments about 800 nm in length and 17 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8542 nt (NC_012038) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Adams et al. 2004).

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

Helminthotheca echioides (Bristly oxtongue)

1147

ToRSV infection in plants of Helleborus foetidus was reported from Lithuania (Navalinskiene et al. 2003). The virus-infected helleborus plants exhibit ringspot symptoms. The virus is transmitted by nematode vectors in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and also by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Cardin L, Onesto JP, Moury B (2003) First report of Cucumber mosaic virus in Helleborus foetidus in France and Italy. Plant Dis 87:1263 Eastwell KC, du Toit LJ, Druffel KL (2009) Helleborus net necrosis virus: a new Carlavirus associated with black death Helleborus sp. Plant Dis 93:332–338 Gera A, Beckelman E, Katzir P, Yam Y, Spiegel S, Zeidan M (2011) New and emerging viruses in ornamental crops. Acta Hortic 901:105–112 Kleinhempel H (1991a) The causal agent of the helleborus ringspot disease. Arch Phytopathol Pflanzenschutz 27:415 Kleinhempel K (1991b) Virus infection of outdoor ornamental plants. In: Report for 1990. Inst Fur Phytopathologie, Aschersleben, p 36 Koenig R (1985) Recently discovered virus or virus-like diseases of ornamentals and their epidemiological significance. Acta Hortic 164:21–31 Liefting LW, Perez-Egusquiza Z, Clover GRG (2010) First report of Helleborus net necrosis virus in Hellebore in New Zealand. Plant Dis 94:479 Murant AF, Roberts JM (1977) An isolate of Broad bean wilt virus serotype II from Helleborus vesicarius. In: Report of Scottish Horticultural Research Institute for 1976. Dundee, Scotland. p 83 Navalinskiene M, Samuitiene M, Jomantiene R (2003) Identification of Tomato ringspot nepovirus and subgroup 16SrI-A of phytoplasmas infecting hellebore plants in Lithuania. Sodininkyste Ir Darzininkyste (Lithuania) 22:259–268 Richert-Poggeler KR, Turhal A-K, Schuhmann S, Maaß C, Blockus S, Zimmermann E, Eastwell KC, Martin RR, Lockhart B (2015) Carlavirus biodiversity in horticultural host plants: efficient virus detection and identification combining electron microscopy and molecular biology tools. Acta Hortic 1072:37–45 Shiraishi T, Hoshi H, Eimori K, Kawanishi T, Komatsu K, Hashimoto M, Maejima K, Yamaji Y, Namba S (2011) First report of hellebore ‘black death’ disease associated with Helleborus net necrosis virus in Japan. J Gen Plant Pathol 77:269–272

Helminthotheca echioides (Bristly oxtongue) Synonyms Helmintia echioides; Picris echioides Family: Asteraceae

Weed host

Pelargonium zonate spot virus Taxonomic position Genus: Anulavirus

(PZSV)

Family: Bromoviridae

PZSV infection in plants of Picris echioides was reported from Italy (Gallitelli et al. 2005). Thrips are involved in transmitting this virus by carrying virus-infected pollen grains on their bodies and introducing the virus into susceptible hosts while feeding. The virus is mechanically sap-transmissible to a wide range of test plants. For more details of PZSV, refer to Pelargonium spp.

H

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Hemerocallis spp. (Daylily)

References Gallitelli D, Finetti-Sialer M, Martelli GP (2005) Anulavirus, a proposed new genus of plant viruses in the family Bromoviridae. Arch Virol 150:407–411

Hemerocallis spp. (Daylily) Family: Xanthorrhoeaceae

Ornamental

Tomato mosaic virus

(ToMV)

Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

ToMV infection in plants of Hemerocallis spp. was reported from Brazil (Duarte et al. 2007). The virusinfected daylily plants show necrotic spots on the foliage. No vector is involved in the spread of this virus, which virus is transmissible by mechanical sap-inoculation, by grafting, and by contact between plants. For more details of ToMV, refer to Solanum lycopersicum.

References Duarte LML, Alexandre MAV, Rivas EB, Cattai MB, Soares RM, Harakava R, Fernandes FMC (2007) Phylogenetic analysis of Tomato mosaic virus from Hemerocallis sp. and Impatiens hawkeri. Summa Phytopathol 33:409–413

Hemidesmus indicus (Indian sarsaparilla) Family: Apocynaceae

Medicinal

Hemidesmus yellow mosaic virus Taxonomic position Genus: Begomovirus

(HemYMV)

Family: Geminiviridae

Geographical distribution HemYMV infection in plants of Hemidesmus indicus was reported from India (Sreekanth Reddy et al. 2014). Symptoms and host(s) The virus-infected Indian sarsaparilla plants exhibit chlorosis, yellow mosaic, and leaf deformation symptoms.

Heracleum spp. (Heracleum sphondylium, H. moellendorfii) (Hogweed)

1149

Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate), incomplete icosahedra, T = 1, 22  38 nm, with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2825 nt (KC898543 = NC_022073) (Briddon 2001; Sreekanth Reddy et al. 2014; Brown et al. 2015; Zerbini et al. 2017).

References Briddon RW (2001) Begomovirus. Geminiviridae. The Springer Index of Viruses. Springer, New York, pp 340–348, https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Sreekanth Reddy M, Kanakala S, Srinivas KP, Hema M, Malathi VG, Sreenivasulu P (2014) Complete genome sequence of a new begomovirus associated with yellow mosaic disease of Hemidesmus indicus in India. Arch Virol 159(5):1223–1228 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Heracleum spp. (Heracleum sphondylium, H. moellendorfii) (Hogweed) Family: Apiaceae

Weed host

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

CGMMV infection in plants of Heracleum moellendorfii was reported from Korea (Cho et al. 2015). The virus-infected hogweed plants exhibit leaf mosaic, mottling, and deformed symptoms. The virus is not transmitted by any insect vector. The virus is mechanically sap-transmissible and also through contact between plants. The virus is transmissible through soil and irrigation water contaminated with infected plant debris. The virus is also pollen transmitted. For more details of CGMMV, refer to Cucumis sativus.

Heracleum latent virus Taxonomic position Genus: Vitivirus

(HLV)

Family: Betaflexiviridae

H

1150

Hesperis matronalis (Dame’s rocket)

Geographical distribution HLV infection was first reported in plants of Heracleum sphondylium from the UK by Murant and Goold (1972). The virus spreads in the Netherlands and the UK (Bem and Murant 1979a). Symptoms and host(s) The virus-infected hogweed plants do not exhibit any symptoms. Transmission The virus is transmitted by aphid vectors Cavariella aegopodii, C. pastinacae, and C. theobaldi in a semipersistent manner. The virus is lost by the vector when it molts, and does not multiply in the vector. The virus is mechanically sap-transmissible and has several experimental hosts in the Amaranthaceae, Chenopodiaceae, Solanaceae, and Umbelliferae. The virus is transmissible by grafting. The virus is not transmissible by contact between plants and by true seed (Bem and Murant 1979a). Virion properties and genome The virions are helically constructed flexuous filaments 730  12 nm, showing distinct crossbanding. The genome is a single molecule of positive-sense ssRNA c. 7.6 kb; a 30 partial genome of 3006 nt is available (X79270). The genome contains five slightly overlapping ORFs which encode, in order, the replication-related proteins, a 19–20 K protein with unknown functions, a MP of the “30K” superfamily type, the CP, and a small protein (10–14 K) with nucleotide-binding properties. Coat protein subunits are of one type and 18–21.5 kDa in size (Bem and Murant 1979b; Adams et al. 2004).

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Bem F, Murant AF (1979a) Transmission and differentiation of six viruses infecting hogweed (Heracleum sphondylium) in Scotland. Ann Appl Biol 92:237–242 Bem F, Murant AF (1979b) Host range, purification and serological properties of heracleum latent virus. Ann Appl Biol 92:243–256 Cho S-Y, Kim YS, Jeon YH (2015) First report of Cucumber green mottle mosaic virus infecting Heracleum moellendorfii in Korea. Plant Dis 99:897 Murant AF, Goold RA (1972) Viruses of umbelliferous plants. Rep Scott Hortic Res Inst 1971:63–64

Hesperis matronalis (Dame’s rocket) Family: Brassicaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Hesperis matronalis was reported from British Columbia (Ford et al. 1988). The virus-infected dame’s rocket plants exhibit symptoms of severe mosaic, distortion, and occasional

Heuchera spp.

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necrosis. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Ribgrass mosaic virus Taxonomic position Genus: Tobamovirus

(RMV)

Family: Virgaviridae

RMV infection in plants of Hesperis matronalis was reported from British Columbia (Ford et al. 1988). The virus-infected dame’s rocket plants exhibit symptoms of severe mosaic, distortion, and occasional necrosis. No vector is reported for this virus. The virus is mechanically sap-transmissible. The virus is transmissible by grafting and also by contact between plants. For more details of RMV, refer to Plantago lanceolata.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Hesperis matronalis was reported from British Columbia (Ford et al. 1988). The virus-infected dame’s rocket plants displayed stunting and leaf curling symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of TuMV, refer to Brassica rapa.

References Ford RE, Beczner L, Hamilton RI (1988) Turnip, cucumber and ribgrass mosaic viruses isolated from Hesperis matronalis in British Columbia. Plant Dis 72:101–106

Heuchera spp. Family: Saxifragaceae

Ornamental

Tobacco rattle virus

(TRV)

Taxonomic position Genus: Tobravirus

Family: Virgaviridae

TRV infection in plants of Heuchera spp. was reported from the USA (Lockhart 2000). The virus-infected heuchera plants exhibit symptoms of yellow ringspotting and concentric line patterns. The virus is transmitted by nematode vectors and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

H

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Hevea brasiliensis (Rubber tree)

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV was reported in plants of Heuchera sanguina in Hungary (Horvath et al. 2006) and probably occurs elsewhere. The virus-infected heuchera plants exhibit leaf deformation and necrotic lesion symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Heuchera spp. was reported from commercial nurseries in the USA (Lockhart et al. 2002). The virus-infected heuchera plants exhibit yellow mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of TuMV, refer to Brassica rapa.

References Horvath J, Baracsi E, Takács A, Kazinczi G, Gáborjányi R, Krajczinger R (2006) Virus infection of ornamental plants in Hungary. Cereal Res Commun 34:485–488 Lockhart BEL (2000) Dicentra, Epimedium, and Heuchera: new perennial ornamental hosts of tobacco rattle virus in the United States. Plant Dis 84:1344 Lockhart BEL, Fetzer J, Westendrop J (2002) Previously unreported viral diseases of Aster, Heuchera, Lobelia, Pulmonaria and Physostegia in the USA. Acta Hortic 568:221–224

Hevea brasiliensis (Rubber tree) Family: Euphorbiaceae

Commercial crop

Tapping Panel Dryness syndrome

(Viroid)

Synonyms Rubber viroid India TPD (tapping panel dryness) syndrome infection in trees of H. brasiliensis was first reported from Malaysia (Sharples 1936). This disease spreads in some South Asian countries including India (Ramachandran et al. 2000; Kumar et al. 2013, 2015).

Hibbertia spp.

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TPD-affected rubber trees show symptoms of bark scaling, cracking, drying, necrotic streaking, and browning of internal bark leading to the DK of internal tissues. Often prominent abnormal bulges on the lower part of tree trunks occur where the first panel begins to dry and leads to reduced latex yield. A viroid with a 360 or 361 nt circular ssRNA genome (NC_014129; 361 nt) was identified in rubber trees with the disease (Ramachandran et al., 2000; Kumar et al., 2015).

References Kumar A, Pandey DM, Abraham T, Mathew J, Ramachandran P, Malathi VG (2013) Determination of biotic aetiology of tapping panel dryness (TPD) syndrome of rubber tree (Hevea brasiliensis) by return-polyacrylamide gel electrophoresis (R-PAGE) technique. Arch Phytopathol Plant Protect 46(6):710–720 Kumar A, Pandey DM, Abraham T, Mathew J, Jyothsna P, Ramachandran P, Malathi VG (2015) Molecular characterization of viroid associated with tapping panel dryness syndrome of Hevea brasiliensis from India. Curr Sci 108(8):1520–1527 Ramachandran P, Mathur S, Francis L, Varma A, Mathew J, Mathew NM, Sethuraj MR (2000) Evidence for association of a viroid with tapping panel dryness syndrome of rubber (Hevea brasiliensis). Plant Dis 84:1155 Sharples S (1936) Brown bast. In: Diseases and pests of the rubber tree. Macmillan, London, pp 229–265

Hewittia spp. Family: Convolvulaceae

Sweet potato feathery mottle virus Taxonomic position Genus: Potyvirus

(SPFMV)

Family: Potyviridae

SPFMV infection in plants of Hewittia spp. was reported from Uganda (Tugume et al. 2010). The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of SPFMV, refer to Ipomoea batatas.

References Tugume AK, Mukasa SB, Valkonen JPT (2010) The significance of wild plants in the evolutionary diversification of Sweet potato feathery mottle virus in East Africa. In: 11th International Plant Virus Epidemiology Symposium held at Cornell University, New York, Abstract

Hibbertia spp. Family: Dilleniaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

H

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Hibiscus cannabinus (Kenaf) (Mesta)

CMV infection in plants of Hibbertia spp. was reported from California (USA) (Endo 1961; Flasinski et al. 1995). The virus-infected hibbertia plants exhibit symptoms of mosaic, chlorotic ringspots, blotches, line patterns, and some leaf deformation. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Hibbertia virus Y Taxonomic position Genus: Potyvirus

(HiVY)

Family: Potyviridae

Geographical distribution HiVY infection in plants of Hibbertia scandens was reported from Australia (Gibbs et al. 2008). Symptoms and host(s) The virus-infected hibbertia plants showing mottled leaves. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense, single-stranded RNA. A partial genome sequence of 1707 nt is available (AF228516) (Revers and Garcia 2015; Wylie et al. 2017).

References Endo RM (1961) A mosaic disease of Hibbertia volubilis. Phytopathology 51:402–406 Flasinski S, Scott SW, Barnett OW, Sun C (1995) Diseases of Peperomia, Impatiens and Hibbertia caused by Cucumber mosaic virus. Plant Dis 79:843–848 Gibbs AJ, Mackenzie AM, Wei K-J, Gibbs MJ (2008) The potyviruses of Australia. Arch Virol 153:1411–1420 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Hibiscus cannabinus (Kenaf) (Mesta) Family: Malvaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Fiber crop

(AMV)

Family: Bromoviridae

Hibiscus cannabinus (Kenaf) (Mesta)

1155

AMV infection in plants of Hibiscus cannabinus was reported from North and South America (RubiesAutonell and Turina 1994). The virus-infected kenaf plants exhibit clear yellow mottle or mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Cotton leaf curl Multan virus Taxonomic position Genus: Begomovirus

(CLCuMuV)

Family: Geminiviridae

CLCuMuV infection in plants of Hibiscus cannabinus was reported from China (Tang et al. 2015). The virus-infected kenaf plants exhibit upwards leaf curling, vein swelling, and dark green vein symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of CLCuMuV, refer to Gossypium spp.

H Hibiscus latent ringspot virus Taxonomic position Genus: Nepovirus

(HLRSV)

Family: Secoviridae

HLRSV infection in plants of Hibiscus cannabinus was reported from Australia, Nigeria, France, Italy, and the USA (Rubies-Autonell and Turina 1995). The virus-infected kenaf plants were stunted, and the leaves were mildly chlorotic. The virus is mechanically sap-transmissible to a number of test plants (Rubies-Autonell and Turina 1995). The intensive studies carried out on seed transmission of HLRSV in kenaf seed, collected from the USA, Australia, and France, indicated up to 26% transmission (Rubies-Autonell and Turina 1997). For more details of HLRSV, refer to Hibiscus rosa-sinensis.

Malvastrum yellow vein virus

(MaYVV)

Synonyms Kenaf leaf curl virus (KLCuV) Taxonomic position Genus: Begomovirus

Family: Geminiviridae

KLCuV infection in plants of Hibiscus cannabinus was reported from India (Paul et al. 2009a). The virus-infected kenaf plants exhibit symptoms of upward leaf curling and stunted growth. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of MaYVV, refer to Malvastrum spp.

Mesta yellow vein mosaic Bahraich virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

(MeYVMBaV)

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Hibiscus cannabinus (Kenaf) (Mesta)

Geographical distribution MeYVMBaV infection in plants of Hibiscus cannabinus was reported from north India (Das et al. 2008; Roy et al. 2009).

Symptoms and host(s) The virus-infected mesta plants show yellowing of veins which leads to complete yellowing of leaves followed by defoliation. Initially the discoloration is greenish yellow at the first stage which develops into yellow mosaic, chlorotic yellowing, and complete chlorosis of foliage. Occasionally the stems of the diseased plants become yellow, partially or totally, and the flowers and fruits are malformed causing low seed yield. Transmission The virus is not mechanically sap-transmissible but is transmitted by the whitefly vector, Bemisia tabaci, with acquisition and inoculation access periods of 12 h each (Das et al. 2008). It is likely that, in common with other begomoviruses, the mechanism of transmission of MeYVMBaV by B. tabaci is in a circulative, non-propagative manner. Experimentally the virus was insect transmitted to H. sabdariffa but not to Nicotiana tabacum, Gossypium hirsutum, or Abelmoschus esculentus. Virion properties and genome The structure of the virions of MeYVMBaV has not been investigated. In common with all geminiviruses, the virions of MeYVMBaV are likely geminate (twinned quasi-icosahedra). MeYVMBaV is a typical Old World monopartite begomovirus. The genome of MeYVMBaV consists of a single circular molecule of a single-stranded DNA of 2737 nt (EU360303 = NC_010818; FJ159267, FJ159268) (Briddon 2001; Das et al. 2008; Roy et al. 2009; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes of MeYVMBaV encode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes of MeYVMBaV have not been investigated.

Mesta yellow vein mosaic virus Taxonomic position Genus: Begomovirus

(MeYVMV)

Family: Geminiviridae

MeYVMV infection in plants of Hibiscus cannabinus was reported from India (Chatterjee et al. 2005; Chatterjee and Ghosh 2007; Ghosh et al. 2007; Rishi 2009). The virus-affected kenaf plants show yellowing of veins and veinlets, which leads to complete yellowing of leaves followed by defoliation. Initially the discoloration is greenish yellow, which develops into yellow mosaic, chlorotic yellowing, and complete chlorosis of foliage in a sequential manner (Chatterjee and Ghosh 2007). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. The virus is graft-transmissible (Chatterjee et al. 2008). For more details of MeYVMV, refer to Hibiscus sabdariffa.

Hibiscus cannabinus (Kenaf) (Mesta)

Tobacco streak virus Taxonomic position Genus: Ilarvirus

1157

(TSV)

Family: Bromoviridae

TSV infection in plants of Hibiscus cannabinus was reported from India and Brazil (Bhaskara Reddy et al. 2012). The virus-infected kenaf plants exhibit symptoms of mosaic, necrotic spotting on leaves, and necrosis of buds. The virus is transmitted by thrips vectors Frankliniella occidentalis and Thrips tabaci; the virus present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is mechanically sap-transmissible. The virus is transmissible by pollen and also by grafting. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato leaf curl Joydebpur virus Taxonomic position Genus: Begomovirus

(ToLCJoV)

H Family: Geminiviridae

ToLCJoV infection in plants of Hibiscus cannabinus was reported from southern India (Paul et al. 2009b). The virus-infected kenaf plants exhibit symptoms of upward curling of leaves with reduction in plant height. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative nonpropagative manner. The virus is graft-transmissible. For more details of ToLCJoV, refer to Solanum lycopersicum.

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Hibiscus cannabinus was reported from India (Raj et al. 2007). The virus-infected kenaf plants exhibit symptoms of yellow leaf vein netting accompanied by excessive yellowing and curling of leaves and stunting of the plants. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is graft-transmissible. For more details of ToLCNDV, refer to Solanum lycopersicum.

References Bhaskara Reddy BV, Sivaprasad Y, Naresh Kumar CVM, Sujitha A, Raja Reddy K, Sai Gopal DVR (2012) First report of Tobacco streak virus infecting Kenaf (Hibiscus cannabinus) in India. Indian J Virol 23(1):80–82 Briddon RW (2001) Begomovirus. Geminiviridae. The Springer Index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Chatterjee A, Ghosh SK (2007) A new monopartite begomovirus isolated from Hibiscus cannabinus L. in India. Arch Virol 152(11):2113–2118 Chatterjee A, Roy A, Padmalatha KV, Malathi VG, Ghosh SK (2005) Occurrence of a Begomovirus with yellow vein mosaic disease of mesta (Hibiscus cannabinus and Hibiscus sabdariffa). Australas Plant Pathol 34:609–610

1158

Hibiscus rosa-sinensis (China rose)

Chatterjee A, Roy A, Ghosh SK (2008) Acquisition, transmission and host range of a begomovirus associated with yellow vein mosaic disease of mesta (Hibiscus cannabinus and Hibiscus sabdariffa). Australas Plant Pathol 37:511–519 Das S, Ghosh R, Paul S, Roy A, Ghosk SK (2008) Complete nucleotide sequence of a monopartite begomovirus associated with yellow vein mosaic disease of mesta from North India. Arch Virol 153:1791–1796 Ghosh R, Paul S, Roy A, Mir JI, Ghosh SK, Srivastava RK, Yadav US (2007) Occurrence of begomovirus associated with yellow vein mosaic disease of kenaf (Hibiscus cannabinus) in northern India. Online. Plant Health Prog. https://doi. org/10.1094/PHP-2007-0508-01-RS Paul S, Ghosh R, Chaudhuri S, Ghosh SK, Roy A (2009a) Biological and molecular variability of the begomoviruses associated with leaf curl disease of kenaf in India. J Plant Pathol 91:637–647 Paul S, Ghosh R, Das S, Palit P, Acharyya S, Das A, Mir JI, Chaudhuri S, Ghosh SK, Roy A (2009b) First report of Tomato leaf curl Joydebpur virus and associated betasatellite in kenaf (Hibiscus cannabinus) plants showing leaf curl symptoms from southern India. Plant Pathol 58:403 Raj SK, Khan MS, Snehi SK, Roy RK (2007) Yellow vein netting of Bimili jute (Hibiscus cannabinus L.) in India caused by a strain of Tomato leaf curl New Delhi virus containing DNA b. Aust Plant Dis Notes 2:45–47 Rishi N (2009) Significant plant virus diseases in India and a glimpse of modern disease management technology. J Gen Plant Pathol 75:1–18 Roy A, Acharyya S, Das S, Ghosh R, Paul S, Srivastava RK, Ghosh SK (2009) Distribution, epidemiology and molecular variability of the begomovirus complexes associated with yellow vein mosaic disease of mesta in India. Virus Res 141(2):237–246 Rubies-Autonell C, Turina M (1994) Alfalfa mosaic virus (AMV) isolated from kenaf (Hibiscus cannabinus). Phytopathol Mediterr 33:234–239 Rubies-Autonell C, Turina M (1995) Characterization and cytopathology of Hibiscus latent ringspot virus isolated from kenaf (Hibiscus cannabinus L.) in Italy. J Phytopathol 143:211–215 Rubies-Autonell C, Turina M (1997) Seed transmission of Hibiscus latent ringspot virus (HLRSV). Plant Dis 81:1082–1084 Tang Y-F, He Z-F, Du Z-G, She X-M, Lan G-B (2015) Detection and identification of the pathogen causing kenaf (Hibiscus cannabinus) leaf curl disease in Hainan Province of China. Acta Phytopathol Sin 45(6):561–568 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Hibiscus rosa-sinensis (China rose) Family: Malvaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Ornamental

(AMV)

Family: Bromoviridae

AMV infection in plants of Hibiscus rosa-sinensis was reported from Spain (Parrella et al. 2012). The virus-infected China rose plants exhibit bright yellow “Aucuba”-type mosaic symptoms. The virus is transmitted by a large number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Citrus leprosis virus C Taxonomic position Genus: Cilevirus

(CiLV-C)

Family: Kitaviridae

Hibiscus rosa-sinensis (China rose)

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CiLV-C was observed to infect plants of Hibiscus rosa-sinensis in Hawaii (Melzer et al. 2013). The virus-infected China rose plants exhibited green ringspot symptoms in senescing leaves. The virus is transmitted by mite vectors (Brevipalpus spp.). The virus is also mechanically sap-transmissible. For more details of CiLV-C, refer to Citrus spp.

Cotton leaf curl Kokhran virus

(CLCuKoV)

Synonyms Cotton leaf curl Burewala virus (CLCuBuV) Taxonomic position Genus: Begomovirus

Family: Geminiviridae

CLCuBuV infection in plants of Hibiscus rosa-sinensis was reported from Pakistan (Akhtar et al. 2014). The virus-infected China rose plants exhibit vein thickening and greening, leaf curling, enations, and stunting symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. The virus is not transmissible by mechanical inoculation, nor by contact between plants. For more details of CLCuKoV, refer to Gossypium spp.

Cotton leaf curl Multan virus Taxonomic position Genus: Begomovirus

(CLCuMuV)

Family: Geminiviridae

CLCuMuV-infected plants of Hibiscus rosa-sinensis was reported from China, the Philippines, and India (Rajeshwari et al. 2005; Mao et al. 2008; Snehi et al. 2013; Gu et al. 2015; Zhang et al. 2015; Srivastava et al. 2016; She et al. 2017). The virus-infected China rose plants exhibit vein thickening, upward curling of leaves and enations on the abaxial leaf surface, reduction in leaf size, and stunting. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. For more details of CLCuMuV, refer to Gossypium spp.

Eggplant mottled dwarf nucleorhabdovirus

(EMDV)

Synonyms Pelargonium vein clearing virus; Pittosporum vein yellowing virus (PVYV); Tomato vein-yellowing virus (TVYV) Taxonomic position Genus: Nucleorhabdovirus

Family: Rhabdoviridae

EMDV infection in plants of Hibiscus rosa-sinensis was reported from the Canary Islands, Yugoslavia, Morocco, Spain, Italy, and Greece (Plavsic et al. 1984; Lockhart 1987; Lawson 1995; De Stradis et al. 2007, 2008; Parrella et al. 2013). The virus-infected China rose plants exhibited vein yellowing and also yellowing of the calyx resulting in failure of flower buds to open fully and premature flower drop. The virus is transmitted by leafhopper vectors in a persistent-propagative manner. The virus is mechanically

H

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Hibiscus rosa-sinensis (China rose)

transmissible from Hibiscus to N. benthamiana, N. rustica, and N. debneyi which produce local and systemic chlorotic spotting and vein yellowing (Plavsic et al. 1984, 1985). For more details of EMDV, refer to Solanum melongena.

Hibiscus chlorotic ringspot virus Taxonomic position Genus: Betacarmovirus

(HCRSV)

Family: Tombusviridae

Geographical distribution HCRSV infection in plants of Hibiscus rosa-sinensis spreads in Iran, Australia, El Salvador, Fiji, Italy, the Solomon Islands, Thailand, Taiwan, New Zealand, Nigeria, Singapore, Malaysia, Bangladesh, and the USA (Jones and Behncken 1980; Kashiwazaki et al. 1982; Raju 1985; Wong and Chang 1992; Abdul-Samad and Mat 1995; Wong et al. 1996; Li and Chang 2002; Tang et al. 2008; Pourrahim et al. 2013; Haque et al. 2014; Tomassoli et al. 2015). Symptoms and host(s) Under natural conditions, China rose plants exhibit mottling or chlorotic spots, rings, or vein-banding symptoms, which may disappear during summer months (Jones and Behncken 1980). Infected hibiscus hybrids have shown severe stunting and flower distortion (Wong and Chang 1992). In addition to H. rosa-sinensis, HCRSV has also been reported to infect H. borianus, and H. densonii (Tomassoli et al. 2015). Transmission The virus is transmissible by mechanical inoculation. This virus incites systemic mosaics in species of Malvaceae. A few members of the Amaranthaceae, Chenopodiaceae, Leguminosae, and Scrophulariaceae respond with local lesions only (Pourrahim et al. 2013). Among the insect vectors tested, aphids like Myzus persicae and beetles like Epilachna varivestis failed to transmit this disease. The main source of virus spread is through planting infected stem cuttings and also through air layering (Waterworth et al. 1976). Raju (1985) has reported the virus transmission through the use of pruning shears. Virion properties and genome The virions are isometric, 30 nm in diameter, and have T = 3 icosahedral symmetry. The genome consists of a single molecule of linear, positive-sense ssRNA of 3911 nt (X86448) containing 7 open reading frames (ORFs) which encode proteins of p28, p81, p8, p9, p38, p23, and p25. In addition to the 3.91 kb genomic RNA, two 30 -co-terminated subgenomic RNAs of approximately 1.7 and 1.5 kb are identified (Huang et al. 2000; Cheng et al. 2009; Luria et al. 2013; Pourrahim et al. 2013).

Hibiscus chlorotic spot virus

(HibCSV)

Taxonomic position HibCSV is a tentative member of the genus Dichorhavirus and family Rhabdoviridae. Geographical distribution HibCSV was reported affecting plants of Hibiscus rosa-sinensis in Brazil (Kitajima et al. 2003).

Hibiscus rosa-sinensis (China rose)

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Symptoms and host(s) The virus-infected China rose plants exhibit chlorotic spot symptoms. Transmission The virus is transmitted by a mite vector, Brevipalpus phoenicis. Virion properties and genome The virions are rod-shaped, 40–50 nm wide by 100–110 nm long, and occur in the nucleus.

Hibiscus green spot virus

(HGSV)

Taxonomic position HGSV is a tentative member of the genus Higrevirus and family Kitaviridae. Geographical distribution HGSV infection in plants of Hibiscus rosa-sinensis was reported from Piracicaba, in Brazilian State of Sao Paulo (Kitajima et al. 2003). Symptoms and host(s) The virus-infected China rose plants exhibit in the tissues green or brown spots or green ringspots in senescent leaves. Transmission The virus is transmitted by a mite vector, Brevipalpus phoenicis (Kitajima et al. 2010). Virion properties and genome The virions are c.60–70  120–150 nm, found in the cytoplasm and occasionally in the perinuclear space, but not internal to the nucleus. No genome information is currently available.

Hibiscus green spot virus 2 Taxonomic position Genus: Higrevirus

(HGSV2)

Family: Kitaviridae

Geographical distribution HGSV2 was reported in plants of Hibiscus arnottianus from Hawaii (Melzer et al. 2012). Hibiscus green spot disease is observed mainly in Hibiscus rosa-sinensis and was initially described from Brazil but has also been observed elsewhere in the Americas, Hawaii, and New Zealand (Melzer et al. 2012). The virus in Brazil is similar, but may be distinct, as it was not detected by primers to HGSV 2. Symptoms and host(s) Leaves of affected China rose plants showed green or brownish spots or ringspots in senescent leaves; symptoms are localized rather than systemic, at sites of mite-transmitted infection. A single tree of Citrus volkameriana growing nearby was also infected, showing symptoms in leaves and bark similar to symptoms of citrus leprosis caused by Citrus leprosis virus-C.

H

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Hibiscus rosa-sinensis (China rose)

Transmission The virus is transmitted by mites of the genus Brevipalpus (Brevipalpus phoenicis) (Roy et al. 2015). Virion properties and genome The virions are short, bacilliform particles (c.30  50 nm). The genome is three molecules of a positivesense, single-stranded RNA: RNA1 consists of 8372 nt (HQ852052 = NC_016141), RNA2 of 3196 nt (HQ852053 = NC_016143), and RNA3 of 3134 nt (HQ852054 = NC_016142).

Hibiscus latent Fort Pierce virus Taxonomic position Genus: Tobamovirus

(HLFPV)

Family: Virgaviridae

Geographical distribution HLFPV infection in plants of Hibiscus rosa-sinensis was observed in different regions of the USA, Italy, and Japan (Adkins et al. 2003; Allen et al. 2005; Yoshida et al. 2014; Tomassoli et al. 2015). Symptoms and host(s) The virus-infected China rose plants exhibit symptoms including diffuse chlorotic spots and rings and an overall chlorotic mottle on the leaves (Allen et al. 2005). In addition to H. rosa-sinensis, the virus has also been reported to infect H. borianus, H. storkly, and H. densonii (Tomassoli et al. 2015). Transmission There is no known vector for this virus. The virus is mechanically sap-transmissible to Hibiscus spp., and also produced local lesions on Chenopodium amaranticolor and Chenopodium quinoa. The virus is easily transmissible through slash and cut inoculation methods (Kamenova and Adkins 2004). The virus spreads by using cuttings from the infected plants for planting. The virus is also transmissible by contact between plants. Virion properties and genome The virions are rigid, rod-shaped particles with dimensions of 15 nm in width and 250–300 nm in length. The genome consists of a positive-sense, single-stranded RNA of 6431 nt (AB917427 = NC_025381) (Allen et al. 2005; Yoshida et al. 2014). The 30 -terminus has a tRNA-like structure. The 50 terminus has a methylated nucleotide cap (m7G50 pppG) (Adams et al. 2017).

Hibiscus latent ringspot virus Taxonomic position Genus: Nepovirus

(HLRSV)

Family: Secoviridae

Geographical distribution HLRSV infection in plants of Hibiscus rosa-sinensis spreads in Australia, the USA, Italy, Bangladesh, and Nigeria (Brunt et al. 1980; Haque et al. 2014).

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Symptoms and host(s) The symptoms on China rose plants are occasional leaf chlorosis, but plants are usually symptomless. Symptoms disappear soon after infection. The infected plants were stunted and did not flower or produce seeds under conditions in northern Italy. Transmission The virus is mechanically sap-transmissible and infects 22 of 74 species from 7 of 20 families when sap inoculated (Brunt et al. 1980). On sap-inoculation, Chenopodium amaranticolour and C. quinoa plants showed systemic flecking symptoms 1 week after inoculation. The virus is transmissible by grafting but not transmissible by contact between plants. Commercial seed lots of various cultivars of kenaf (Hibiscus cannabinus) were shown to transmit HLRSV to progeny seedlings in different percentages up to a maximum of 26% (Rubies-Autonell and Turina 1997), but not transmitted in limited tests with seed from Nicotiana clevelandii and Chenopodium quinoa. Transmission by a nematode vector is suspected. Virion properties and genome The virions are isometric, non-enveloped of two types, but of similar size, 25–30 nm in diameter, and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is a bipartite, linear, positive-sense, single-stranded RNA (Sanfacon et al. 2009; Dunez and Le Gall 2011; Sanfacon 2015; Thompson et al. 2017); no sequence is currently available.

Hibiscus latent Singapore virus

(HLSV)

Synonyms Hibiscus virus S Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

Geographical distribution A tobamovirus infecting plants of Hibiscus rosa-sinensis in Singapore was shown to be distinct and initially named Hibiscus virus S (Srinivasan et al. 2002, 2005) but later renamed Hibiscus latent Singapore virus. Isolates of HLSV have also been identified in Taiwan (Chen et al. 2001; Liu et al. 2004) and Italy (Tomassoli et al. 2015). Symptoms and host(s) The virus-infected China rose plants exhibit vein yellowing and mosaic symptoms. Transmission Transmission occurs without the help of vectors. The virus is mechanically sap-transmissible and also through contact between plants. Virion properties and genome The virions are non-enveloped, rigid helical rods with a helical symmetry, about 18 nm in diameter, and 300–310 nm in length. The genome is a monopartite, linear, positive-sense ssRNA of 6485 nt (AF395898 = NC_008310). The 30 -terminus has a tRNA-like structure. The 50 terminus has a

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methylated nucleotide cap (m7G50 pppG) (Srinivasan et al. 2005; Zaitlin 2011; Niu et al. 2014; Adams et al. 2017).

Hop stunt viroid

(HpSVd)

Taxonomic position Genus: Hostuviroid

Family: Pospiviroidae

HpSVd isolate was detected in plants of Hibiscus rosa-sinensis in Italy (Sanger 1988; Luigi et al. 2013). HpSVd-infected China rose plants showed both severe symptoms of reduction in plant growth and upward curling and deformation of leaves, during surveys made in Italy by Luigi et al. (2013). HpSVd has a very wide host range (Sanger 1988). Since the occurrence of HpSVd, infected Hibiscus rosasinensis is very frequent, this host may act as a source of inoculum for infection of economically important fruit crops. The viroid is transmissible by mechanical sap-inoculation, and through vegetative propagation and grafting. For more details of HpSVd, refer to Humulus lupulus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV was reported in plants of Hibiscus rosa-sinensis in New Zealand (Elliott et al. 2009), without specific symptom descriptions. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Okra mosaic virus Taxonomic position Genus: Tymovirus

(OkMV)

Family: Tymoviridae

OkMV infection in plants of Hibiscus rosa-sinensis was reported from the Ivory Coast and Nigeria (Givord et al. 1972; Igwegbe 1983). The virus-infected China rose plants exhibit mild mosaic symptoms. The virus is transmitted by beetles of the genus Podagrica in a semi-persistent manner (Lana and Taylor 1975). The virus is mechanically sap-transmissible and also by grafting. For more details of OkMV, refer to Abelmoschus esculentus.

Tomato mosaic virus Taxonomic position Genus: Tobamovirus

(ToMV)

Family: Virgaviridae

ToMV infection in plants of Hibiscus rosa-sinensis was reported from China (Huang et al. 2004). The systemic symptoms induced by this virus were dark and light green mosaic in young leaves, leaf

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puckering and malformation on older leaves, and significant stunting. No vector is involved in the spread of this virus. This virus is transmissible by mechanical sap-inoculation, by grafting, and by contact between plants. For more details of ToMV, refer to Solanum lycopersicum.

References Abdul-Samad N, Mat M (1995) Hibiscus chlorotic ringspot virus in Malaysian hibiscus. Plant Dis 79:967 Adams MJ, Adkins S, Bragard C, Gilmer D, Li D, MacFarlane SA, Wong S-M, Melcher U, Ratti C, Ryu KH, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Virgaviridae. J Gen Virol 98:1999–2000 Adkins S, Kamenova I, Achor D, Lewandowski DJ (2003) Biological and molecular characterization of a novel Tobamovirus with a unique host range. Plant Dis 87:1190–1196 Akhtar KP, Ullah R, Saeed M, Sarwar N, Mansoor S (2014) China rose (Hibiscus rosa-sinensis): a new natural host of Cotton leaf curl Burewala virus in Pakistan. J Plant Pathol 96:385–389 Allen JE, Kamenova I, Adkins S, Hanson SF (2005) First report of Hibiscus latent Fort Pierce virus in New Mexico. Plant Health Prog. https://doi.org/10.1094/PHP-2005-0105-01-HN Brunt AA, Barton RJ, Phillips S, Lana O (1980) Hibiscus latent ringspot virus: a newly recognised virus from Hibiscus rosa-sinensis (Malvaceae) in western Nigeria. Ann Appl Biol 96:37–43 Chen TH, Kao KH, Liou MR, Kao CL (2001) A unique tobamovirus-like virus isolated from vein yellowing mosaic China rose (Hibiscus rosa-sinensis L.) in Taiwan. Plant Pathol Bull 10:195–200 Cheng A, Speir JA, Yuan YA, Johnson JE, Wong SM (2009) Preliminary X-ray data analysis of crystalline Hibiscus chlorotic ringspot virus. Acta Crystallogr Sect F: Struct Biol Cryst Commun 65:589–593 De Stradis A, Parrella G, Vovlas C, Ragozzino A (2007) A rhabdovirus associated with hibiscus vein clearing disease in Southern Italy. J Plant Pathol 89(3):S39 De Stradis A, Parrella G, Vovlas C, Ragozzino A (2008) Vein yellowing of Hibiscus rosa-sinensis caused by Eggplant mottled dwarf virus in southern Italy. J Plant Pathol 90:359–361 Dunez J, Le Gall O (2011) Nepovirus. Comoviridae. In: The Springer Index of Viruses. Springer, New York, pp 361–369. https://doi.org/10.1007/978-0-387-95919-1_55 Elliott DR, Lebas BSM, Ochoa-Corona FM, Tang J, Alexander BJR (2009) Investigation of Impatiens necrotic spot virus outbreaks in New Zealand. Australas Plant Pathol 38:490–495 Givord L, Pfeiffer P, Hirth L (1972) Un nouveau virus du groupe de la Mosaique Jaune du Navet: le virus de la Mosaique du Gombo. CR Acad Sci Paris Ser D 275:1563–1566 Gu Z-H, Hu G-J, Xie Y, Zhou X-P (2015) Construction and pathogenicity evaluation of infectious clones of Cotton leaf curl Multan virus infecting Hibiscus rosa-sinensis and Gossypium hirsutum [J]. Acta Phytopathol Sin 45(2):198–204 Haque MAS, Akanda AM, Sultana R, Majumdar A, Rahman MA (2014) An investigation to virus like disease of China rose. Bangladesh Res Pub J 10(2):125–137 Huang M, Koh DCY, Weng L-J, Chang M-L, Yap Y-K, Zhang L, Wong S-M (2000) Complete nucleotide sequence and genome organization of Hibiscus chlorotic ringspot virus, a new member of the genus Carmovirus: evidence for the presence and expression of two novel open reading frames. J Virol 74:3149–3155 Huang JG, Fan ZF, Li HF, Tian GZ, Hu JS (2004) First report of Tomato mosaic virus on Hibiscus rosa-sinensis in China. Plant Dis 88:683 Igwegbe ECK (1983) New strain of Okra mosaic virus in Nigeria. Plant Dis 67:320–322 Jones DR, Behncken GM (1980) Hibiscus chlorotic ringspot, a widespread virus disease in the ornamental Hibiscus rosa–sinensis. Aust Plant Pathol 9:4–5 Kamenova I, Adkins S (2004) Transmission, in planta distribution and management of Hibiscus latent fort pierce virus, a novel Tobamovirus isolated from Florida hibiscus. Plant Dis 88:674–679 Kashiwazaki T, Yamashita S, Doi Y (1982) Two hibiscus-infecting viruses: Hibiscus yellow mosaic virus and Hibiscus chlorotic ringspot virus. Ann Phytopathol Soc Jpn 48:395 Kitajima EW, Chagas CM, Rodrigues JCV (2003) Brevipalpus-transmitted plant virus and virus-like diseases: cytopathology and some recent cases. Exp Appl Acarol 30:135–160 Kitajima EW, Rodrigues JCV, Freitas-Astúa J (2010) An annotated list of ornamentals naturally found infected by Brevipalpus mite transmitted viruses. Sci Agric 67:348–371 Lana AO, Taylor TA (1975) The insect transmission of an isolate of okra mosaic virus occurring in Nigeria. Ann Appl Biol 82:361–364 Lawson RH (1995) Hibiscus. In: Loebenstein G, Lawson RH, Brunt AA (eds) Virus and virus-like diseases of bulb and flower crops. Wiley, Chichester, pp 476–481 Li S-C, Chang Y-C (2002) First report of Hibiscus chlorotic ringspot virus in Taiwan. Plant Pathol 51(6):803

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Liu FL, Chen TC, Yeh SD, Hsu HT, Chen CC, Bau HJ, Chen YK (2004) Serological and molecular characterizations of a hibiscus-infecting tobamovirus in Taiwan. Plant Pathol Bull 13:283–290 Lockhart BEL (1987) Evidence for identity of plant rhabdoviruses causing vein-yellowing diseases of tomato and Hibiscus rosa-sinensis. Plant Dis 71:731–733 Luigi M, Manglli A, Tomassoli L, Faggioli F (2013) First report of Hop stunt viroid in Hibiscus rosa-sinensis in Italy. New Dis Rep 27:14 Luria N, Reingold V, Lachman O, Dombrovsky A (2013) Full-genome sequence of Hibiscus chlorotic ringspot virus from Israel. Genome Announc 1(6):e01050-13. https://doi.org/10.1128/genomeA.01050-13 Mao MJ, He ZF, Yu H, Li HP (2008) Molecular characterization of Cotton leaf curl Multan virus and its satellite DNA that infects Hibiscus rosa-sinensis. Chin J Virol 24:64–68 Melzer MJ, Sether DM, Borth WB, Hu JS (2012) Characterization of a virus infecting Citrus volkameriana with citrus leprosis-like symptoms. Phytopathology 102:122–127 Melzer MJ, Simbajon N, Carillo J, Borth WB, Freitas-Astúa J, Kitajima EW, Neupane KR, Hu JS (2013) A cilevirus infects ornamental hibiscus in Hawaii. Arch Virol 158:2421–2424 Niu S, Cao S, Wong SM (2014) An infectious RNA with a hepta-adenosine stretch responsible for programmed -1 ribosomal frameshift derived from a full-length cDNA clone of Hibiscus latent Singapore virus. Virology 449:229–234 Parrella G, Fiallo-Olivé E, Navas-Castillo J (2012) First report of China rose (Hibiscus rosa-sinensis) as a host of Alfalfa mosaic virus in Spain. Plant Dis 96:462 Parrella G, Ade S, Greco B, Villanueva F, Fortes IM, Navas-Castillo J (2013) First report of Eggplant mottled dwarf virus in China rose in southern Spain. Span J Agric Res 11(1):204–207 Plavsic B, Erič Z, Milicič D (1984) Rhabdovirus-like particles associated with vein yellowing of Hibiscus rosa-sinensis L. Phytopathol Mediterr 23:52–54 Plavsic B, Milicic D, Eric Z (1985) Rhabdovirus – like particles in Hibiscus rosa-sinensis L. Acta Hortic 164:41–44 Pourrahim R, Ghobakhlo A, Farzadfar S (2013) Biological and molecular detection of Hibiscus chlorotic ringspot virus infecting Hibiscus rosa-sinensis in Iran. Phytopathol Mediterr 52:528–531 Rajeshwari R, Reddy RVC, Maruthi MN, Colvin J, Seal SE, Muniyappa V (2005) Host range, vector relationships and sequence comparison of a begomovirus infecting hibiscus in India. Ann Appl Biol 147:15–25 Raju BC (1985) Occurrence of chlorotic ringspot virus in commercial Hibiscus rosa-sinensis cultivars. Acta Hortic 164:273–280 Roy A, Hartung JS, Schneider WL, Shao J, León MG, Melzer MJ, Otero-Colina G, Beard JJ, Bauchan GR, Ochoa R, Brlansky RH (2015) Role bending: complex relationships between viruses, hosts and mite vectors related to citrus leprosis, an emerging disease. Phytopathology 105:1013–1025 Rubies-Autonell C, Turina M (1997) Seed transmission of Hibiscus latent ringspot virus (HLRSV). Plant Dis 81:1082–1084 Sanfacon H (2015) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. In: Encyclopedia of Life Sciences. Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000764.pub3. http:// www.els.net Sanfacon H, Wellink J, Le Gall O, Karasev A, van der Vlugt R, Wetzel T (2009) Secoviridae: a proposed family of plant viruses within the order Picornavirales that combines the families Sequiviridae and Comoviridae, the unassigned genera Cheravirus and Sadwavirus, and the proposed genus Torradovirus. Arch Virol 154(5):899–907 Sanger HL (1988) Viroids and viroid diseases. Acta Hortic 234:79–87 She XM, Tang YF, He ZF, Lan GB (2017) Molecular characterization of Cotton leaf curl Multan virus and its associated betasatellite infecting Hibiscus rosa-sinensis in the Philippines. J Plant Pathol 99(3):765–768 Snehi SK, Srivastava A, Kumar S, Raj SK (2013) Molecular detection and identification of begomoviruses affecting important ornamental plants in India. Indian J Virol 24:142 Srinivasan KG, Narendrakumar R, Wong SM (2002) Hibiscus virus S is a new subgroup II tobamovirus: evidence from its unique coat protein and movement protein sequences. Arch Virol 147:1585–1598 Srinivasan KG, Min BE, Ryu KH, Adkins S, Wong SM (2005) Determination of complete nucleotide sequence of Hibiscus latent Singapore virus: evidence for the presence of an internal poly(A) tract. Arch Virol 150:153–166 Srivastava A, Kumar S, Jaidi M, Raj SK (2016) Association of Cotton leaf curl Multan virus and its associated betasatellite with leaf curl disease of Hibiscus rosa-sinensis in India. New Dis Rep 33:4 Tang J, Elliott DR, Quinn BD, Clover GRG, Alexander BJR (2008) Occurrence of Hibiscus chlorotic ringspot virus in Hibiscus spp. in New Zealand. Plant Dis 92:1367 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J Gen Virol 98:529–531 Tomassoli L, Manglli A, Tiberini A, Adkins S (2015) Investigation on the phytosanitary status of major ornamental Hibiscus species in Italy to assess virus infection. Acta Hortic 1072:29–35

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Waterworth HE, Lawson RH, Monroe RL (1976) Purification and properties of Hibiscus chlorotic ring spot virus. Phytopathology 66:570–575 Wong SM, Chang CG (1992) Occurrence of Hibiscus chlorotic ringspot virus in Singapore. Phytopathology 82:722 Wong SM, Chng CG, Chong PL, Lee YH (1996) Characterization of a Singapore isolate of hibiscus chlorotic ringspot virus. Acta Hortic 432:76–83 Yoshida T, Kitazawa Y, Komatsu K, Neriya Y, Ishikawa K, Fujita N, Hashimoto M, Maejima K, Yamaji Y, Namba S (2014) Complete nucleotide sequence and genome structure of a Japanese isolate of Hibiscus latent Fort Pierce virus, a unique tobamovirus that contains an internal poly(A) region in its 30 end. Arch Virol 159:3161–3165 Zaitlin M (2011) Tobamovirus. In: Tidona C, Darai G (eds) The Springer Index of Viruses. Springer, New York. https:// doi.org/10.1007/978-0-387-95919-1 Zhang H, Ji Y-H, Wu S-H, Zhao W-H, Zhou T, Zhou Y-J (2015) Identification and characterization of Cotton leaf curl Multan virus from Hibiscus rosa-sinensis in Jiangsu. Acta Phytopathol Sin 45(4):361–369

Hibiscus sabdariffa (Roselle) (Mesta) Family: Malvaceae

Mesta yellow vein mosaic virus Taxonomic position Genus: Begomovirus

H

Fiber crop

(MeYVMV)

Family: Geminiviridae

Geographical distribution MeYVMV infection in plants of Hibiscus sabdariffa was reported from India (Chatterjee et al. 2005; Roy et al. 2009). Symptoms and host(s) The virus-affected mesta plants show yellowing of veins and veinlets, which leads to complete yellowing of leaves, followed by defoliation. Initially the discoloration is greenish yellow, processing to yellow mosaic, chlorotic yellowing, and complete chlorosis of foliage in a sequential manner. Occasionally the stems of the diseased plants become yellow, partially or totally, and the flowers and fruits are malformed causing low seed yield. The virus has been isolated from mesta (Hibiscus cannabinus and Hibiscus sabdariffa), Malvastrum coromandelianum, and hollyhock (Alcea rosea). Transmission The virus is transmitted by the whitefly vector Bemisia tabaci with an acquisition and inoculation access period of 12 h each. It is likely that, in common with other begomoviruses, the mechanism of transmission of MeYVMV by B. tabaci is in a circulative, non-propagative manner. The virus is not mechanically sap or seed-transmitted. The virus is graft-transmissible (Chatterjee et al. 2005, 2008). Virion properties and genome The structure of the virions of MeYVMV has not been investigated. In common with all geminiviruses, the virions of MeYVMV are likely geminate (twinned quasi-icosahedra). MeYVMV is a typical Old World monopartite begomovirus. The genome of MeYVMV consists of a single circular molecule of a single-stranded DNA 2752 nt (EF373060 = NC_009088; FJ159269;

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EF428256) (Briddon 2001; Roy et al. 2009; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes of MeYVMVencode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes of MeYVMV have not been investigated. Some isolates of the virus have been shown to associate with a betasatellite (Das et al. 2008).

Okra mosaic virus Taxonomic position Genus: Tymovirus

(OkMV)

Family: Tymoviridae

OkMV infection in plants of Hibiscus sabdariffa was reported from Ivory Coast (Givord 1978). The virus-infected roselle plants exhibit chlorotic blotches. The virus is transmitted by beetle vectors in a semi-persistent manner; by mechanical sap-inoculation, and by grafting. For more details of OkMV, refer to Abelmoschus esculentus.

References Briddon RW (2001) Begomovirus. Geminiviridae. The Springer Index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Chatterjee A, Roy A, Padmalatha KV, Malathi VG, Ghosh SK (2005) Occurrence of a Begomovirus with yellow vein mosaic disease of mesta (Hibiscus cannabinus and Hibiscus sabdariffa). Australas Plant Pathol 34:609–610 Chatterjee A, Roy A, Ghosh SK (2008) Acquisition, transmission and host range of a begomovirus associated with yellow vein mosaic disease of mesta (Hibiscus cannabinus and Hibiscus sabdariffa). Australas Plant Pathol 37:511–519 Das S, Roy A, Ghosh R, Paul S, Acharyya S, Ghosh S (2008) Sequence variability and phylogenetic relationship of betasatellite isolates associated with yellow vein mosaic disease of mesta in India. Virus Genes 37(3):414–424 Givord L (1978) Alternative hosts of Okra mosaic virus near plantings of okra in Southern Ivory Coast. Plant Dis Reptr 62:412–416 Roy A, Acharyya S, Das S, Ghosh R, Paul S, Srivastava RK, Ghosh SK (2009) Distribution, epidemiology and molecular variability of the begomovirus complexes associated with yellow vein mosaic disease of mesta in India. Virus Res 141:237–246 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Hibiscus syriacus (Rose of sharon) Family: Malvaceae

Ornamental

Eggplant mottled dwarf nucleorhabdovirus Taxonomic position Genus: Nucleorhabdovirus

(EMDV)

Family: Rhabdoviridae

EMDV infection in plants of Hibiscus syriacus was reported from Australia (Tang et al. 2015). The virus-infected rose of Sharon plants exhibit symptoms of chlorotic spots on leaves. The virus is transmitted by leafhopper vectors in a persistent-propagative manner. The virus is transmissible by

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mechanical sap-inoculation. The virus spreads primarily through the use of infected planting material. For more details of EMDV, refer to Solanum melongena.

Euphorbia yellow leaf curl virus Taxonomic position Genus: Begomovirus

(EuYLCV)

Family: Geminiviridae

EuYLCV infection in plants of Hibiscus syriacus was reported from Pakistan (Riaz et al. 2015). The virus-infected Rose of Sharon plants show leaf curling and yellowing symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. For more details of EuYLCV, refer to Euphorbia pulcherrima.

H Reference Riaz H, Ashfaq M, Mukhtar T, Riaz T (2015) First report of Euphorbia yellow leaf curl virus infecting Hibiscus syriacus. J Plant Pathol 97:S67 Tang J, Elliott C, Ward LI, Iqram WA (2015) Identification of Eggplant mottled dwarf virus in PEQ Hibiscus syriacus plants imported from Australia. Australas Plant Dis Notes 10:6

Hippeastrum spp. (Amaryllis) Family: Amarilidaceae

Ornamental

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Hippeastrum spp. was reported from South Africa (Schulze et al. 2017). The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Capsicum chlorosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(CaCV)

Family: Tospoviridae

CaCV infection in plants of Hippeastrum spp. was reported from Taiwan (Chen et al. 2009, 2012). The virus-infected amaryllis plants exhibit chlorotic ringspot symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of CaCV, refer to Capsicum annuum.

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Cucumber mosaic virus

(CMV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV infection in plants of Hippeastrum hybridum was reported from Mexico and India (GutierrezVillegas et al. 2004; Adhab and Al-Ani 2011). The virus-infected amaryllis plants show yellow concentric rings and line patterns on the leaves. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. The virus can also be transmissible by using contaminated knives when cutting the flowers (Kahn and Scott 1964; Loebenstein et al. 1995). For more details of CMV, refer to Cucumis sativus.

Hippeastrum chlorotic ringspot virus

(HCRV)

Taxonomic position HCRV is a tentative member of the genus Orthotospovirus and family Tospoviridae Geographical distribution HCRV was reported from plants of Hippeastrum spp. in China (Dong et al. 2013; Xu et al. 2014). Symptoms and host(s) The virus-infected plants of amaryllis showed necrotic and chlorotic ringspot symptoms on the foliage. Infections of Hymenocallis littoralis, Clivia miniata and Zephyranthes candidahave also been reported from China (e.g. HG763861, KY484838, and KY363498, respectively). Transmission The virus is transmitted by thrips vectors, Thrips palmi and T. tabaci, in a persistent-propagative manner. The virus is also mechanically sap-transmissible. Virion properties and genome Quasi-spherical enveloped virions of 70–100 nm were observed in tissue extracts and thin sections. The genome is comprised of three RNAs, the L, M, and sRNA segments, of 8908 nt (L RNA; HG763861, KY363498, and KY484838), 4762-4764 nt (M RNA; KY363497 and KY484837), and 2744-2749 nt (KC290943, KY363496, and KY484836) respectively (Xu et al. 2014).

Hippeastrum latent virus Taxonomic position Genus: Carlavirus

Family: Betaflexiviridae

Geographical distribution The virus infection in plants of Hippeastrum hybridum was reported from the Netherlands and Taiwan (Brolman-Hupkes 1975; Wang and Tai unpublished - NC_011540). Symptoms and host(s) The virus-infected amaryllis plants do not exhibit any obvious symptoms.

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Transmission The virus is mechanically sap-transmissible and also through infected planting material. Virion properties and genome The virions are flexuous filaments about 584–611 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8500 nt (NC_011540, DQ098905) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Adams et al. 2004).

Hippeastrum mosaic virus Taxonomic position Genus: Potyvirus

(HiMV)

Family: Potyviridae

H

Geographical distribution HiMV infection in plants of Hippeastrum hybridium was first reported from the Netherlands (Kunkel 1922). The virus spreads in Australia, Brazil, the former Czechoslovakia, Fiji, Greece, India, Japan, the Netherlands, South Africa, the UK, China, and the USA (Brants and van den Heuvel 1965; Raj et al. 2009; Alexandre et al. 2011; Pham et al. 2011; Wylie and Jones 2012; Malandraki et al. 2016; Xu et al. 2017). Symptoms and host(s) The virus-infected amaryllis leaves and flower stalks show an irregular light and dark green mosaic patterns. Transmission The virus is transmitted by aphid vectors, Aphis fabae, A. gossypii, and Myzus persicae, in a nonpersistent manner. The virus is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants, not transmissible by seed, and not transmissible by pollen. Virion properties and genome The virions are filaments, non-enveloped, usually flexuous, and with a clear modal length of 782 nm and 12 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9660 nt (JQ395040 = NC_017967) (Wylie and Jones 2012; Revers and Garcia 2015; Wylie et al. 2017).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Hippeastrum hortorum was reported from the Czech Republic (Mertelik et al. 2002). The virus-infected amaryllis plants exhibit chlorotic spots and line pattern symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

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Hippeastrum spp. (Amaryllis)

Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

IYSV has been detected in plants of amaryllis (Hippeastrum hybridum “Orange Sovereign”) in Israel (Gera et al. 1998; Kritzman et al. 2001, 2008). The virus-infected amaryllis plants showed chlorotic spots and rings (Kritzman et al. 2001). The virus is transmitted by the onion thrips vector, Thrips tabaci, in a persistent-propagative manner, and is also transmissible by the mechanical sap-inoculation. For more details of IYSV, refer to Iris spp. or Allium cepa.

Nerine latent virus Taxonomic position Genus: Carlavirus

(NeLV)

Family: Betaflexiviridae

NeLV infection in plants of Hippeastrum spp. was reported from Australia and Singapore (Wong et al. 1996; Wylie and Jones 2012). The virus-infected amaryllis plants exhibit chlorotic striping symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of NeLV, refer to Nerine spp.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV infection in plants of Hippeastrum hybridum was reported from the Netherlands (De Leeuw 1972). The virus-infected amaryllis plants exhibit mosaic symptoms. There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Hippeastrum spp. was reported from Poland and the Netherlands (Kaminska and Korbin 1994; Derks 1995; Derks and Lemmers 1996). The virus-infected amaryllis plants showed yellowish foliar spots and red necrotic lesions, with leaves finally turning yellow and dying prematurely but without affecting flowering. Virus-free seedlings of hippeastrum infected by thrips transmission in a greenhouse developed irregular-shaped white or dark green spots and line patterns. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Hippeastrum spp. (Amaryllis)

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References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Adhab MA, Al-Ani RA (2011) Amaryllis and shrimp plant are secondary hosts of Cucumber mosaic cucumovirus (CMV) in Iraq. Agric Biol J N Am 2:872–875 Alexandre MAV, Duarte LML, Rivas EB, Cilli A, Harakava R, Galleti SR, Kitajima EW (2011) Hippeastrum mosaic virus diagnosed in hippeastrum and eucharis in Brazil. J Plant Pathol 93:643–649 Brants DH, van den Heuvel J (1965) Investigation of Hippeastrum mosaic virus in Hippeastrum hybridum. Neth J Plant Pathol 71:145–151 Brolman-Hupkes JE (1975) Tentative description of Hippeastrum latent virus in Hippeastrum hybridum plants and differentiation from Hippeastrum mosaic virus. Neth J Plant Pathol 81:226–236 Chen CC, Huang CH, Cheng YH, Chen TC, Yeh SD, Chang CA (2009) First report of Capsicum chlorosis virus infecting Amaryllis and Blood Lily in Taiwan. Plant Dis 93:1346 Chen C-C, Chang C-A, Yeh S-D, Chen T-C (2012) Occurrence of Capsicum chlorosis virus on ornamental crops in Taiwan. Bioforsk Fokus 7(9):42 De Leeuw GTN (1972) Tobacco mosaic virus in Hippeastrum hybridum. Neth J Plant Pathol 78:69–71 Derks AFLM (1995) Hippeastrum. In: Loebenstein G, Lawson RH, Brunt AA (eds) Virus and virus-like diseases of bulb and flowering crops. Wiley, UK, pp 293–297 Derks AFLM, Lemmers MEC (1996) Detection of tospoviruses in bulbous crops and transmissibility by vegetative propagation. Acta Hortic 432:132–139 Dong JH, Yin YY, Fang Q, McBeath JH, Zhang ZK (2013) A new tospovirus causing chlorotic ringspot on Hippeastrum sp. in China. Virus Genes 46:567–570 Gera A, Kritzman A, Cohen J, Raccah B (1998) Tospoviruses infecting bulb crops in Israel. In: Peters D, Goldbach R (eds) Recent progress in tospovirus and thrips research. Abstracts of Papers and Poster Presentations Presented at the Fourth International Symposium on Tospoviruses and Thrips in Floral and Vegetable Crops, held 2–6 May 1998 in Wageningen, The Netherlands. pp 86–87 Gutierrez-Villegas C, Ruiz-Medrano R, Piedra-Ibarra E, de la Torre-Almaraz R (2004) Characterization of a Cucumber mosaic virus strain associated with yellow mottle symptoms of amaryllis (Hippeastrum hybridum Leopoldii) in Mexico. Agrociencia (Montecillo) 38:343–354 Kahn RP, Scott HA (1964) Serological relationship of Cucumber mosaic virus and certain virus isolates that incite Amaryllis mosaic symptoms. Phytopathology 54:360–362 Kaminska M, Korbin M (1994) New natural hosts of Tomato spotted wilt virus. Acta Hortic 377:123–128 Kritzman A, Lampel M, Raccah B, Gera A (2001) Distribution and transmission of Iris yellow spot virus. Plant Dis 85:838–842 Kritzman A, Beckelman Y, Tam Y, Raccah B, Gera A (2008) The current status of iris yellow spot virus in Israel. The 3rd Conference of the International Working Group on Legume and Vegetable Viruses (IWGLVV), August 20th–23rd, 2008, p 33 Kunkel LO (1922) Ameboid bodies associated with Hippeastrum mosaic. Science 55:73 Loebenstein G, Lawson RH, Brunt AA (1995) Virus and virus-like diseases of bulb and flowering crops. Wiley, UK, p 543 Malandraki I, Driessen A, Varveri C, Vassilakos N (2016) First report of Hippeastrum mosaic virus in Hippeastrum sp. in Greece. Plant Dis 100:869 Mertelik J, Mokra V, Gotzova B, Gabrielova S (2002) Occurrence and identification of impatiens necrotic spot tospovirus in the Czech Republic. Acta Hortic 568:79–83 Pham KTK, de Kock MJD, Lemmers MEC, Derks AFLM (2011) Molecular identification of potyviruses infecting bulbous ornamentals by the analysis of coat protein (CP) sequences. Acta Hortic 901:167–172 Raj SK, Snehi SK, Kumar S, Khan MS (2009) First molecular detection and identification of a potyvirus associated with severe mosaic disease of amaryllis (Hippeastrum hybridum Hort.) in India. Australas Plant Dis Notes 4:50–53 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Schulze A, Roberts R, Pietersen G (2017) First report of the detection of bean yellow mosaic virus (BYMV) on Tropaeolum majus; Hippeastrum spp., and Liatris spp. in South Africa. Plant Dis 101:846 Wong SM, Chng CG, Chong PL (1996) Identification of nerine latent virus in Hippeastrum in Singapore. Phytoparasitica 24(4):327 Wylie SJ, Jones MGK (2012) Complete genome sequences of seven carlavirus and potyvirus isolates from Narcissus and Hippeastrum plants in Australia, and proposals to clarify their naming. Arch Virol 157:1471–1480 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

H

1174

Hippuris vulgaris (Mare’s tail or Horsetail)

Xu Y, Lou S-G, Li Q-F, Hu Q, Sun L-P, Li Z-Y, Li Y-Z, Liu Y-T (2014) Molecular characterization of the hippeastrum chlorotic ringspot virus L segment and its protein. Arch Virol 159:2805–2807 Xu XH, Tang W, Gao R, Yang SK, Li F, Sun HW, Lu XB (2017) First report of Hippeastrum mosaic virus in Hippeastrum spp. in mainland China. Plant Dis 101:1064

Hippuris vulgaris (Mare’s tail or Horsetail) Family: Plantaginaceae

Medicinal

Alfalfa mosaic virus

(AMV)

Taxonomic position Genus: Alfamovirus

Family: Bromoviridae

AMV infection in plants of Hippuris vulgaris was reported from Saudi Arabia (Al-Shahwan et al. 2017). The virus is transmitted by several species of aphids in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Bean common mosaic virus Taxonomic position Genus: Potyvirus

(BCMV)

Family: Potyviridae

BCMV infection in plants of Hippuris vulgaris was reported from Saudi Arabia (Al-Shahwan et al. 2017). The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BCMV, refer to Phaseolus vulgaris.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Hippuris vulgaris was reported from Saudi Arabia (Al-Shahwan et al. 2017). The virus is transmitted by the thrips vectors; the virus present in/on pollen entering the host through injuries caused by thrips while feeding. The virus is mechanically sap-transmissible and not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Reference Al-Shahwan IM, Abdalla OA, Al-Saleh MA, Amer MA (2017) Detection of new viruses in alfalfa, weeds and cultivated plants growing adjacent to alfalfa fields in Saudi Arabia. Saudi J Biol Sci 24(6):1336–1343

Holcus lanatus (Velvet grass)

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Hirschfeldia incana (Shortpod mustard) Synonyms Brassica geniculata Family: Brassicaceae

Weed host

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Hirschfeldia incana was reported from New Zealand (Fletcher 2001). The virus-infected shortpod mustard plants do not exhibit obvious symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Hirschfeldia incana was reported from Cyprus (Papayiannis et al. 2011). The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. The virus is transmitted by grafting but not transmitted by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

Reference Fletcher JD (2001) New hosts of Alfalfa mosaic virus, Cucumber mosaic virus, Potato virus Y, Soybean dwarf virus, and Tomato spotted wilt virus in New Zealand. N Z J Crop Hortic Sci 29:213–217 Papayiannis L, Katis NI, Idris AM, Brown JK (2011) Identification of weed hosts of Tomato yellow leaf curl virus in Cyprus. Plant Dis 95:120–125

Holcus lanatus (Velvet grass) Family: Poaceae

Forage crop

Holcus lanatus yellowing virus

(HLYV)

Taxonomic position HLYV is a tentative member of the family Rhabdoviridae

H

1176

Hordeum jubatum (Foxtail barley)

Geographical distribution HLYV infection was first reported in plants of Holcus lanatus from Italy by Amici et al. (1978). Symptoms and host(s) The virus-infected velvet grass plants exhibit clear leaf chlorosis symptoms. Transmission There is no known vector for this virus, but an arthropod vector is suspected. Virion properties and genome The virions are bullet-shaped.

Ryegrass mosaic virus

(RGMV)

Taxonomic position Genus: Rymovirus

Family: Potyviridae

RGMV infection in plants of Holcus lanatus was reported from New Zealand (Guy 2006). The virus is transmitted by the mite vector Abacarus hystrix in a semi-persistent manner and also through mechanical sap-inoculation. For more details of RGMV, refer to Lolium spp.

Tomato spotted wilt orthotospovirus Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Holcus lanatus was reported from Italy (Grieco et al. 2000). The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Amici A, Faoro F, Tornaghi R (1978) Natural occurrence of Rhabdovirus-like particles in thin sections of different plant species Avena sativa, Chrysanthemum frutescens, Holcus lanatus, Ranunculus repens, Secale cereal, oats, ornamental plant, rye. Riv Patol Veg 14:85–98 Grieco PD, Conte D, Munno I, Nuzzaci M, de Stradis A (2000) Tomato spotted wilt virus (TSWV) on weeds and wild plants in Metapontino, Basilicata. Informatore Fitopatologico 50:43–46 Guy PL (2006) New Zealand grasslands revisited: identification of Cocksfoot mild mosaic virus. Australas Plant Pathol 35:461–464

Hordeum jubatum (Foxtail barley) Family: Poaceae

Weed host

Hordeum mosaic virus Taxonomic position Genus: Rymovirus

(HoMV)

Family: Potyviridae

Hordeum murinum (False barley)

1177

HoMV infection in plants of Hordeum jubatum was reported from Alberta, Canada (Slykhuis and Bell 1966). The virus-infected foxtail barley plants exhibit leaf mottling symptoms. No vector is known for this virus, but an eriophyid mite vector is suspected. The virus is mechanically sap-transmissible. For more details of HoMV, refer to Hordeum vulgare.

Reference Slykhuis JT, Bell W (1966) Differentiation of agropyron mosaic, wheat streak mosaic, and a hitherto unrecognized hordeum mosaic virus in Canada. Can J Bot 44:1191–1208

Hordeum murinum (False barley) Family: Poaceae

Brome streak mosaic virus Taxonomic position Genus: Tritimovirus

H

Fodder crop

(BrSMV)

Family: Potyviridae

BrSMV infection in plants of Hordeum murinum was reported from Austria (Schubert and Rabenstein 1995; Rabenstein and Huss 2013). The virus-infected false barley plants exhibit chlorotic leaf streaking symptoms. The virus is transmitted by an eriophyid mite vector in a semi-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BrSMV, refer to Bromus spp.

Wheat streak mosaic virus Taxonomic position Genus: Tritimovirus

(WSMV)

Family: Potyviridae

WSMV infection in plants of Hordeum murinum was reported from Germany (Rabenstein et al. 1982). The virus is transmitted by eriophyid mite vectors in a semi-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of WSMV, refer to Triticum aestivum.

References Rabenstein F, Huss H (2013) Studies on grass viruses in Austria. In: 63. Jahrestagung der Vereinigung der Pflanzenzüchter und Saatgutkaufleute Österreichs, Raumberg-Gumpenstein, Österreich, 15–18 Rabenstein F, Stanarius A, Proeseler G (1982) Identification of wheat streak mosaic virus in Hordeum murinum L. in the GDR. Arch Phytopathol Pflanzenschutz 18:301–318 Schubert J, Rabenstein F (1995) Sequence of the 30 -terminal region of the RNA of a mite transmitted Potyvirus from Hordeum murinum L. Eur J Plant Pathol 101:123–132

1178

Hordeum vulgare (Barley)

Hordeum vulgare (Barley) Family: Poaceae

Cereal

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

ArMV infection in plants of Hordeum vulgare was reported from Switzerland (Ramel et al. 1994). The virus-infected barley plants exhibit partial stunting and yellowing symptoms. The virus is transmitted by a nematode vector, Xiphinema diversicaudatum, in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Barley mild mosaic virus Taxonomic position Genus: Bymovirus

(BaMMV)

Family: Potyviridae

Geographical distribution BaMMV was first reported as a strain of Barley yellow mosaic virus (called the M or Streatley isolate). BaMMV infection in plants of Hordeum vulgare has been reported from China, Japan, Korea, Belgium, Germany, France, the Netherlands, Greece, Denmark, Ukraine, Great Britain, and Spain (Fantakhun et al. 1987; Hariri et al. 1990; Huth 1990; Adams 1991; Chen et al. 1992; Dessens and Meyer 1995; Lee et al. 1996; Nomura et al. 1996; Katis et al. 1997; Zheng et al. 1999; Achon et al. 2003; Rabenstein and Huss 2013). Symptoms and host(s) The virus-infected barley fields show yellow patches or appear entirely yellow, depending on the distribution of infested soil. The initial symptoms are irregularly distributed chlorotic streaks of varying size on the youngest emerging leaves, often associated with upward rolling of leaf margins. The streaks are most distinct on the youngest leaves and develop into a mosaic pattern. The mosaic symptoms are occasionally associated with necrotic patches, yellow discoloration, and/or rapid death of the older leaves. The symptoms on leaves appear in early spring but tend to disappear as the weather becomes warmer. The virus has a narrow host range, limited to the Gramineae. Transmission The virus is transmitted by the fungal vector, Polymyxa graminis, in a persistent manner. Zoospores, released either from resting spores (cystosori) or from zoosporangia in barley roots, transmit the virus (Jianping et al. 1991; Peerenboom et al. 1996). The virus is not known to be seed-borne. Virion properties and genome The virions are non-enveloped, flexuous filaments of two modal lengths, 250–300 nm and 500–600 nm; both are 13 nm in width. The genome consists of two molecules of a linear, positive-sense ssRNA.

Hordeum vulgare (Barley)

1179

RNA-1 is of 7262 nt (Y10973 = NC_003483) and RNA-2 of 3524 nt (X90904 = NC_003482) (Timpe and Kuhne 1994; Kashiwazaki 1996; Lee et al. 1996; Peerenboom et al. 1996; Zheng et al. 1999; Lopez-Moya et al. 2009; Adams 2011; Wylie et al. 2017).

Barley stripe mosaic virus Taxonomic position Genus: Hordeivirus

(BSMV)

Family: Virgaviridae

Geographical distribution BSMV infection in plants of Hordeum vulgare is distributed worldwide including the Eurasian region, the North American region, and the Pacific region and Australia, England, France, Germany, Poland, Denmark, Yugoslavia, Asia, the UK, the USA, Yemen, and the former USSR (Carroll 1980; Lister et al. 1981; Zamora et al. 1988; Donald et al. 1993; Najar et al. 2000; Arneodo and Truol 2001; Herrera et al. 2001; Jezewska 2001; Kumari et al. 2006; Smith et al. 2014). Symptoms and host(s) The virus-infected barley plants exhibit symptoms of mild stripe mosaic and yellow or whitish chlorosis that appear first near the base of the youngest leaves. In the chronic systemic infection stage, chlorotic stripes broken by yellow and whitish areas form between the veins of developed leaves. Once these parts and areas merge, the leaves take on a generally chlorotic appearance. At the final stage, infected plants may express lethal necrosis, depending on the virus strain (Bragg et al. 2008). Transmission The virus is transmissible by mechanical sap-inoculation. Seed transmission up to 90–100% is recorded (Hamilton 1965; Carroll 1972; La and Park 1979; Lister et al. 1981), and the virus is also transmissible by pollen to the pollinated plant. The virus can also be transmissible from plant to plant when leaves rub together as a result of wind, hail, or animals. No natural vectors of the virus are known. Virion properties and genome The virions are rod-shaped, non-enveloped, and usually straight, with a clear modal length of 100–150 nm and 18–24 nm wide. The genome consists of three positive-sense, single-stranded RNAs: RNA1 consists of 3768 nt (J04342 = NC_003469), RNA2 of 3289 nt (X03854 = NC_003481), and RNA3 of 3164 nt (M16576 = NC_003478) and a “triple gene block” set of cellto-cell movement proteins (Jackson and Lane 1981; Gustafson and Armour 1986; Gustafson et al. 1987, 1989; Jackson et al. 2011; Adams et al. 2017).

Barley yellow dwarf virus

(BYDV)

Taxonomic position BYDV is a tentative member of the genus Luteovirus and family Luteoviridae BYDV is a name that encompasses several member of the genera Luteovirus and Polerovirus (Family Luteoviridae) and two currently unassigned species of the family Luteoviridae. Barley yellow dwarf virus PAV is the type species of the genus Luteovirus; other species in this genus are Barley yellow dwarf virus PAS, Barley yellow dwarf virus MAV, Barley yellow dwarf virus kerII, and Barley yellow dwarf

H

1180

Hordeum vulgare (Barley)

virus kerIII. Barley yellow dwarf virus GPV and Barley yellow dwarf virus SGV are currently unassigned to a genus. Cereal yellow dwarf virus RPS and Cereal yellow dwarf virus RPV, and Maize yellow dwarf virus RMV are members of the genus Polerovirus. Maize yellow mosaic virus is another species in the genus Polerovirus recognized by ICTV in 2017, and not previously noted under the umbrella of ‘Barley yellow dwarf virus’. Earlier reports, and some more recent reports, do not identify the virus beyond the broad name ‘Barley yellow dwarf virus’. Geographical distribution BYDV infection in plants of Hordeum vulgare was first reported by Oswald and Houston (1951). The virus spreads in California, North America, Europe, Greece, the Czech Republic, Africa, Australia, and Asia (Nagaich and Vashisth 1963; Rochow 1969; Panayotou 1980; Halbert and Pike 1986; El Yamani and Hill 1990; D’Arcy 1995; Lister and Ranieri 1995; Miller and Rosachova 1997). Symptoms and host(s) In barley, BYDV infection causes a characteristic bright yellowing of the leaves (particularly older leaves) and pale yellow stripes between the leaf veins plus chlorotic blotching of young leaves. In some varieties, reddening of leaf tips also develops. Early infected barley plants are stunted and produce low grain yields and shriveled grain. Tillering can be stimulated by infection, but most tillers then develop poorly and produce sterile heads. Plants infected after tillering have milder symptoms, and yields are less severely affected (D’Arcy 1995). The natural hosts of this virus are only monocotyledonous members of the Gramineae family. Transmission Over 25 species of aphids are known to transmit different isolates and species of ‘BYDV’. However, only four are common: the bird cherry-oat aphid (Rhopalosiphum padi), the corn leaf aphid (Rhopalosiphum maidis), the English grain aphid (Macrosiphum avenae), and the greenbug (Schizaphis graminum) (Jedlinski 1981; Zhang et al. 1983; Power and Gray 1995). The aphids transmit this virus in a circulative, non-propagative manner. The virus is retained when the vector molts, but does not multiply in the vector, nor is it transmitted congenitally to progeny of the vector. The virus is not mechanically transmissible nor by contact between plants and is not transmissible by seed and pollen (Makkouk et al. 2001a). Virion properties and genome The virions are isometric, non-enveloped, and 25–28 nm in diameter. The genome consists of a positive-sense, single-stranded RNA (Hammond et al. 1983; Martin and D’Arcy 1995).

Barley yellow dwarf virus GPV Taxonomic position Genus: Unassigned

(BYDV-GPV)

Family: Luteoviridae

Geographical distribution BYDV-GPV infection in plants of Hordeum vulgare was reported from China (Zhang et al. 2009). Symptoms and host(s) The virus-infected barley plants exhibit yellowing or reddening symptoms.

Hordeum vulgare (Barley)

1181

Transmission The virus is transmitted by an aphid vector, Rhopalosiphum padi, in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. Virion properties and genome The virions are isometric. The genome consists of a positive-sense, single-stranded RNA; only a partial sequence of 606 nt of many isolates is available (e.g. EU386825 = NC_039035).

Barley yellow dwarf virus kerII Taxonomic position Genus: Luteovirus

(BYDV-kerII)

Family: Luteoviridae

Geographical distribution BYDV-kerII infection in plants of Hordeum vulgare was reported from subantarctic Kerguelen Islands (Svanella-Dumas et al. 2013). Symptoms and host(s) The virus-infected barley plants exhibit yellowing or reddening symptoms. Transmission The virus is transmitted by the aphid vector, Rhopalosiphum padi, in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. Virion properties and genome The virions are 25–30 nm in diameter, are hexagonal in outline, and have no envelope. The genome is a monopartite, linear, positive-sense, single-stranded RNA of 5736 nt (KC571999 = NC_021481). The 30 terminus has neither a poly(A) tract nor a tRNA-like structure. There is no genome-linked protein (VPg) at the 50 -terminus. The genome consists of five or six ORFs that are predicted to encode proteins between 4 and 132 kDa.

Barley yellow dwarf virus kerIII Taxonomic position Genus: Luteovirus

(BYDV-kerIII)

Family: Luteoviridae

Geographical distribution BYDV-kerIII infection in plants of Hordeum vulgare was reported from subantarctic Kerguelen Islands (Svanella-Dumas et al. 2013). Symptoms and host(s) The virus-infected barley plants exhibit yellowing or reddening symptoms. Transmission The virus is transmitted by the aphid vector, Rhopalosiphum padi, in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible.

H

1182

Hordeum vulgare (Barley)

Virion properties and genome The virions are 25–30 nm in diameter, are hexagonal in outline, and have no envelope. The genome is a monopartite, linear, positive-sense, single-stranded RNA; a partial genome of 4625 nt is available (KC559092). The 30 terminus has neither a poly(A) tract nor a tRNA-like structure. There is no genome-linked protein (VPg) at the 50 -terminus. The genome consists of five or six ORFs that are predicted to encode proteins between 4 and 132 kDa.

Barley yellow dwarf virus MAV Taxonomic position Genus: Luteovirus

(BYDV-MAV)

Family: Luteoviridae

Geographical distribution BYDV-MAV infection in plants of Hordeum vulgare was reported from the USA, the Czech Republic and Tunisia (Rochow, 1969, 1979; Vincent et al. 1990; Makkouk et al. 2001a; Kundu 2009; Najar et al. 2017). Symptoms and host(s) The virus-infected barley plants exhibit yellowing or reddening symptoms. Transmission The virus is transmitted by aphid vectors, Sitobion avenae and Metopolophium dirhodum, in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. Virion properties and genome The virions are 25–30 nm in diameter, are hexagonal in outline, and have no envelope. The genome is a monopartite, linear, positive-sense, single-stranded RNA of 5273 nt (D11028 = NC_003680). The 30 terminus has neither a poly (A) tract nor a tRNA-like structure. There is no genome-linked protein (VPg) at the 50 -terminus. The genome consists of five or six ORFs that are predicted to encode proteins between 4 and 132 kDa (Ueng et al. 1992).

Barley yellow dwarf virus PAS Taxonomic position Genus: Luteovirus

(BYDV-PAS)

Family: Luteoviridae

Geographical distribution BYDV-PAS infection in plants of Hordeum vulgare was reported from the Czech Republic and Poland (Kundu 2008; Trzmiel 2017). Symptoms and host(s) The virus-infected barley plants exhibit yellowing or reddening symptoms. Transmission The virus is transmitted by an aphid vector, Rhopalosiphum padi, in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible.

Hordeum vulgare (Barley)

1183

Virion properties and genome The virions are 25–30 nm in diameter, are hexagonal in outline, and have no envelope. The genome is a monopartite, linear, positive-sense, single-stranded RNA of 5695 nt (AF218798 = NC_002160). The 30 terminus has neither a poly(A) tract nor a tRNA-like structure. There is no genome-linked protein (VPg) at the 50 -terminus. The genome consists of five or six ORFs that are predicted to encode proteins between 4 and 132 kDa.

Barley yellow dwarf virus PAV Taxonomic position Genus: Luteovirus

(BYDV–PAV)

Family: Luteoviridae

Geographical distribution BYDV-PAV infection in plants of Hordeum vulgare was reported from the USA, subantarctic Kerguelen Islands, Australia, Yemen, and Tunisia (Rochow, 1969,1979; Bencharki et al. 2000; Makkouk et al. 2001a; Robertson 2003; Kumari et al. 2006; Ansi et al. 2007; Svanella-Dumas et al. 2013; Najar et al. 2000; Maligate et al. 2016; Najar et al. 2017). Symptoms and host(s) The virus-infected barley plants exhibit yellowing or reddening symptoms. Transmission The virus is transmitted by the aphid vectors, Rhopalosiphum padi and Schizaphis avenae, in a circulative, non-propagative manner (Bencharki et al. 2000; Mirshekar et al. 2013). The virus is not mechanically sap-transmissible. Virion properties and genome The virions are 25–30 nm in diameter, are hexagonal in outline, and have no envelope. The genome is a monopartite, linear, positive-sense, single-stranded RNA of 5677 nt (X07653 = NC_004750). The 30 terminus has neither a poly (A) tract nor a tRNA-like structure. There is no genome-linked protein (VPg) at the 50 -terminus. The genome consists of five or six ORFs that are predicted to encode proteins between 4 and 132 kDa (Ueng et al. 1992).

Barley yellow dwarf virus SGV Taxonomic position Genus: Unassigned

(BYDV–SGV)

Family: Luteoviridae

Geographical distribution BYDV-SGV infection in plants of Hordeum vulgare was reported from Poland, the USA, and Tunisia (Makkouk et al. 2001a; Malmstrom and Shu 2004; Trzmiel 2017). Symptoms and host(s) The virus-infected barley plants exhibit yellowing or reddening symptoms.

H

1184

Hordeum vulgare (Barley)

Transmission The virus is transmitted by an aphid vector, Schizaphis graminum, in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible.

Virion properties and genome The virions are isometric. The genome consists of a positive-sense, single-stranded RNA of c.5.5 kb; partial sequences of up to 1267 nt (AY541038) are available.

Barley yellow mosaic virus Taxonomic position Genus: Bymovirus

(BaYMV)

Family: Potyviridae

Geographical distribution BaYMV infection was first recognized in Japan in 1940. The virus infection in plants of Hordeum vulgare was reported from Europe, Asia, Belgium, Greece, China, France, the Netherlands, Belgium, Germany, Poland, Italy, Japan, Korea, Spain, and the UK (Hill 1980; Hill and Evans 1980; Adams et al. 1986, 1987; Anon 1989; Kashiwazaki et al. 1989; Adams 1991; Katis et al. 1997; Achon et al. 2005; Jezewska and Trzmiel 2009, 2010; Rabenstein and Huss 2013).

Symptoms and host(s) On BaYMV-infected barley plants, under field conditions, symptoms only occur in crops sown in the autumn, and they appear during winter and spring, usually when growth resumes after a spell of cold weather (sub-zero temperatures). In virus-infected barley, the mosaic symptoms are seen on leaves of the individual plants. These are elongated, pale green or yellow flecks, typically on the youngest (unfurling) leaves. On some barley varieties, infected leaves may be curled, giving the plants a spiky appearance, and there may also be chocolate-brown necrosis associated with the lesions. Affected plants have fewer tillers and yield less grain, and the grain size may be irregular (Hill and Evans 1980). The natural hosts of this virus include Hordeum vulgare, H. sativum, and H. spontaneum.

Transmission The virus is transmitted by the fungal vector, Polymyxa graminis, in a persistent manner (Adams et al. 1986, 1988). The virus is mechanically sap-transmissible with difficulty to some species of Hordeum and to a few other species of the Poaceae.

Virion properties and genome The virions are non-enveloped, flexuous filaments of two modal lengths, 250–300 nm and 500–600 nm; both are 13 nm in width. The genome consists of two molecules of a linear, positive-sense ssRNA. RNA-1 is of 7637 nt (AJ132268 = NC_002990) and RNA-2 of 3582 nt (AJ132269 = NC_002991) (Huth et al. 1984; Kashiwazaki et al. 1990, 1991; Davidson et al. 1991; Peerenboom et al. 1992; Chen et al. 1999; Lopez-Moya et al. 2009; Adams 2011; Wylie et al. 2017).

Hordeum vulgare (Barley)

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Barley yellow streak mosaic virus

(BaYSMV)

Taxonomic position BaYSMV is a tentative member of the genus Cytorhabdovirus and family Rhabdoviridae. Geographical distribution BaYSMV infection in plants of Hordeum vulgare was reported from Canada and the USA (Robertson and Carroll 1989, 1991; Skaf et al. 1992; Robertson and Brumfield 2000). Symptoms and host(s) The virus-infected barley plants exhibit symptoms of yellow green mosaic, streaking, color banding, and severe necrosis. The typical symptoms are light green to yellow dashes, streaks, and stripes parallel to the midrib. Transmission The virus is transmitted by a mite vector, Petrobia latens (Smidansky and Carroll 1996). The virus is transmissible mechanically to Echinochloa crus-galli, Hordeum vulgare, Lolium persicum, Setaria viridis, and Triticum aestivum and producing mosaic and banding. Local lesions were produced on Chenopodium quinoa and Nicotiana benthamiana. The virus is not transmissible by contact between plants. No seed and pollen transmissions are observed (Robertson and Carroll 1991). Virion properties and genome The virions are unusually shaped and enveloped, usually flexuous, with no clear modal length of 127–4000 nm and 52–81 nm wide. Basic helix is obscure. The genome consists of a single-stranded RNA (Robertson and Carroll 1988; Skaf and Carroll 1995).

Barley yellow striate mosaic cytorhabdovirus

(BYSMV)

Synonyms Maize sterile stunt virus; Wheat chlorotic streak virus Taxonomic position Genus: Cytorhabdovirus

Family: Rhabdoviridae

Geographical distribution BYSMV infection in plants of Hordeum vulgare was reported from the African region, the Eurasian region, the Middle East, and the Pacific region and Australia, France, Iran, Italy, Syria, Lebanon, Tunisia, and Morocco (Conti 1972, 1988; Lockhart et al. 1986; Najar et al. 2000; Makkouk et al. 1996, 2001b; Makkouk 2003). Symptoms and host(s) The virus-infected barley plants exhibit symptoms of yellow striate mosaic, pronounced stunting and changeable striate mosaic symptoms on leaves, chlorotic striations, leaf striating, and plant apical yellowing. The natural hosts of this virus are only monocots of the family Gramineae including wheat, Agropyron repens, oats, rye, and maize.

H

1186

Hordeum vulgare (Barley)

Transmission The virus is transmitted by the planthopper vectors, Laodelphax striatellus and Javesella pellucida, in a persistent-propagative manner. The virus is not transmissible by either mechanical inoculation or through seed (Conti and Plumb 1977; Conti 1980; Makkouk 2003). Virion properties and genome The virions are bacilliform, with a modal length of 270 nm and 60 nm wide. There are distinct surface projections (spikes) dispersed evenly over all the surface. The genome is a monopartite, negative-sense, single-stranded RNA of 12,706 nt (KM213865 = NC_028244) is unsegmented (Appiano and Conti 1974; Dietzgen 2011; Yan et al. 2015; Walker et al. 2018).

Brome mosaic virus Taxonomic position Genus: Bromovirus

(BMV)

Family: Bromoviridae

BMV infection in plants of Hordeum vulgare was reported from Europe, Australia, Finland, South Africa, and the USA (Bremer 1973; Pocsai 1987; Cooper 1988; Pocsai et al. 1991). The virusinfected barley plants exhibit leaf symptoms which are most obvious on young plants and fade as plants mature, often disappearing by heading. Initially, light yellow or white spots and streaks occur that spread rapidly, giving leaves a bright yellow mosaic pattern. Diseased plants are dwarfed and have shriveled heads. The virus is transmitted by the cereal leaf beetle vector, Oulema melanopus, and is also transmissible by mechanical sap-inoculation to many monocotyledonous and a few dicotyledonous plants. For more details of BMV, refer to Bromus spp.

Brome streak mosaic virus Taxonomic position Genus: Tritimovirus

(BrSMV)

Family: Potyviridae

BrSMV infection in plants of Hordeum vulgare was reported from southern France (Huth et al. 1995). The virus is transmitted by an eriophyid mite, Aceria tulipae, in a semi-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of BrSMV, refer to Bromus spp.

Cereal chlorotic mottle virus

(CCMoV)

Taxonomic position CCMoV is a tentative member of the genus Nucleorhabdovirus and family Rhabdoviridae. Geographical distribution CCMoV infection in plants of Hordeum vulgare was reported from Northern Africa and Australia (Greber 1977, 1979, 1981, 1982; Lockhart 1986). Symptoms and host(s) The virus-infected barley plants show stunting and severe necrotic or chlorotic streaks on the leaves.

Hordeum vulgare (Barley)

1187

The natural host range of this virus is confined to the family Graminae, including Avena sativa, Bromus unioloides, Digitaria ciliaris, Dinebra retroflexa, Echinochloa colona, Eleusine indica, Eragrostis cilianensis, Hordeum vulgare, Leptochloa filiformis, Secale cereale, Setaria italica, Setaria verticillata, Triticum aestivum, T. monococcum, T. tauschii, Urochloa panicoides, and Zea mays. Transmission The virus is efficiently transmitted by at least two species of cicadellid leafhopper vectors, Nesoclutha pallida and Cicadulina bimaculata, in a persistent-propagative manner (Greber 1977, 1979, 1981). The virus usually persists in the vector for life, but some insects cease to transmit up to 4 weeks before death. The virus is acquired both by adults and by nymphs and is transmitted through the molt. The virus infects cells of the brain and salivary glands of both vector species and apparently multiplies in the vector. The virus is not mechanically sap-transmissible. Not transmitted through seed (Lockhart 1986). Virion properties and genome The virions are bacilliform particles of 65  240 nm size particles. The genome is a single-stranded RNA (Greber 1979; Walker et al. 2018), presumed to be of negative sense.

Cereal yellow dwarf virus RPS Taxonomic position Genus: Polerovirus

(CYDV-RPS)

Family: Luteoviridae

Geographical distribution CYDV-RPS infection in plants of Hordeum vulgare was reported from the USA (Beckett and Miller, unpublished - NC_002198). Symptoms and host(s) The virus-infected barley plants exhibit bright yellow mosaic symptoms. Transmission The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. Virion properties and genome The virions are 25–30 nm in diameter, are hexagonal in outline, and have no envelope. The genome is a positive-sense, single-stranded RNA of 5662 nt (AF235168 = NC_002198). The 30 terminus has neither a poly(A) tract nor a tRNA-like structure. The 50 terminus has a genome-linked protein (VPg) (of about 17 kDa). The genome consists of six open reading frames (ORFs).

Cereal yellow dwarf virus RPV Taxonomic position Genus: Polerovirus

(CYDV-RPV)

Family: Luteoviridae

H

1188

Hordeum vulgare (Barley)

Geographical distribution CYDV-RPV infection in plants of Hordeum vulgare was reported from Tunisia, Yemen, and Latvia (Makkouk et al. 2001a; Bisnieks et al. 2006; Kumari et al. 2006; Najar et al. 2017). Symptoms and host(s) The virus-infected barley plants exhibit bright yellow mosaic symptoms. Transmission The virus is transmitted by at least 25 species of aphid vectors in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. Virion properties and genome The virions are 25–30 nm in diameter, are hexagonal in outline, and have no envelope. The genome is a positive-sense, single-stranded RNA of approximately 5723 nt (L25299 = NC_004751) (Vincent et al. 1990). The 30 terminus has neither a poly(A) tract nor a tRNA-like structure. The 50 terminus has a genome-linked protein (VPg) (of about 17 kDa). The genome consists of six open reading frames (ORFs).

Chloris striate mosaic virus

(CSMV)

Synonyms Wheat striate mosaic virus Taxonomic position Genu: Mastrevirus

Family: Geminiviridae

CSMV infection in plants of Hordeum vulgare was reported from Australia (Grylls 1963). The virusinfected barley plants exhibit yellow, broken streaks and dwarfing symptoms. The virus is transmitted by a leafhopper vector, Nesoclutha obscura, in a persistent, circulative, and non-propagative manner. The virus is not seed or sap-transmissible. For more details of CSMV, refer to Chloris gayana.

Hordeum mosaic virus Taxonomic position Genus: Rymovirus

(HoMV)

Family: Potyviridae

Geographical distribution HoMV infection in plants of Hordeum vulgare was reported from Alberta, Canada (Salm et al. 1996; French and Stenger 2005). Symptoms and host(s) The virus-infected barley plants exhibit diffuse chlorotic mottle symptoms. Transmission No vector is identified for this virus, which is assumed to be mite-transmitted. The virus is transmissible by mechanical inoculation but not transmitted by seed and not transmitted by pollen.

Hordeum vulgare (Barley)

1189

Virion properties and genome The virions are non-enveloped, flexuous filaments 701 nm long and 15.5 nm in diameter. The genome consists of a single molecule of linear, positive-sense ssRNA of 9463 nt (AY623627 = NC_005904) with a 30 -poly(A) terminus (Salm et al. 1996; Zagula et al. 1992; Lopez-Moya et al. 2009; French and Stenger 2005; Wylie et al. 2017).

Hordeum vulgare alphaendornavirus Taxonomic position Genus: Alphaendornavirus

(HvEV)

Family: Endornaviridae

Geographical distribution HvEV infection in plants of Hordeum vulgare was reported from Europe (Candresse et al. 2016). Symptoms and host(s) The virus-infected barley plants do not exhibit any external symptoms. Transmission The virus is transmitted through seed via both ova and pollen. No horizontal spread has been observed in the field, and no potential vectors have been identified. The virus is not mechanically sap-transmissible. Virion properties and genome The virus does not have a gene for a capsid protein; therefore no virion is formed. The genome is a double-stranded RNA of 14,243 bp, with 50 and 30 noncoding regions of 207 and 47 bp, respectively (Candresse et al. 2016). The monocistronic genome encodes a single large protein of 4663 amino acids and contains three conserved domains: a MTR, a Hel-1, and an RdRp (KT721705).

Indian peanut clump virus Taxonomic position Genus: Pecluvirus

(IPCV)

Family: Virgaviridae

IPCV infection in plants of Hordeum vulgare was reported from India (Delfosse et al. 1999). The virus-infected barley plants were stunted and bushy, with chlorotic or necrotic leaves; most of the infected plants died. The virus is transmitted by a fungal vector and is also transmissible by mechanical sap-inoculation. For more details of IPCV, refer to Arachis hypogaea.

Maize streak virus

(MSV)

Synonyms African cereal streak virus Taxonomic position Genus: Mastrevirus

Family: Geminiviridae

H

1190

Hordeum vulgare (Barley)

MSV infection in plants of Hordeum vulgare was reported from East Africa, Ethiopia, and Kenya (Harder and Bakker 1973; Harder 1975; Shepherd et al. 2010). The virus-infected barley plants initially produce faint, broken, chlorotic streaks which begin near the leaf base and extend upward. Eventually, well-defined alternate yellow and green streaks develop along the entire leaf blade, which eventually becomes completely chlorotic. Leaves formed after infection develop a “shoestring” habit and die. Plants infected while young remain severely stunted and die prematurely; phloem necrosis may be observed. The inflorescence becomes yellow, and awned types in particular become distorted. The virus is transmitted by the leafhopper vectors, Cicadulina mbila, C. arachidis, C. bipunctella, C. triangular, C. bimaculata, C. similes, C. lateens, C. ghaurii, and C. parazeae, in a persistent, circulative, and nonpropagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of MSV, refer to Zea mays.

Mal de Rio Cuarto virus Taxonomic position Genus: Fijivirus

(MRCV)

Family: Reoviridae

MRCV infection in plants of Hordeum vulgare was reported from Argentina (Arneodo and Truol 2001). The virus is transmitted in a propagative way by the planthopper Delphacodes kuscheli (Arneodo et al. 2002). For more details of MRCV, refer to Zea mays.

Northern cereal mosaic cytorhabdovirus

(NCMV)

Synonyms Wheat rosette stunt virus Taxonomic position Genus: Cytorhabdovirus

Family: Rhabdoviridae

Geographical distribution NCMV infection in plants of Hordeum vulgare was first reported from Japan by Ito and Fukushi (1944). The virus spreads in China, Japan, Siberia, Korea, and Pakistan (Shikata and Lu 1967; Hyung and Shikata 1977; Ogawa and Moichi 1984). Symptoms and host(s) The virus-infected barley plants exhibit primary symptoms of infection consisting of chlorotic spots on the basal parts of elongating leaves. The chlorotic spots coalesce and develop into chlorotic stripe or mosaic symptoms. The mosaic symptoms differ slightly with plant species; they usually fade as the plant ages and temperatures rise. Leaves developing later are short and narrow and show chlorotic stripes or generalized yellowing. The growth of infected plants is much decreased, and chlorotic tillers develop one by one, resulting in a stunted rosette appearance. The natural hosts of this virus are barley, Italian millet, oat, rye, Italian ryegrass, wheat, Digitaria violascens, Poa annua, Setaria pycrocoma, and S. viridis.

Hordeum vulgare (Barley)

1191

Transmission The virus is transmitted by planthopper vectors, Laodelphax striatellus, Ribantodelphax albifascia, Unkanodes albifascia, U. sapporona, and Muellerianella fairmairei, in a persistent-propagative manner. The virus is reported not to be transmitted through eggs to the progeny of the vector. The virus is not transmissible by either mechanical sap-inoculation or by contact between plants. The virus is not transmitted through seed (Ruan et al. 1983). Virion properties and genome The virions are enveloped, bacilliform, and measures 60 nm in diameter and 300–350 nm long. There are distinct surface projections (spikes) dispersed evenly over all the surface. The genome is a monopartite, negative-sense, single-stranded RNA of 13,222 nt (AB030277 = NC_002251) is unsegmented (Tanno et al. 2000; Dietzgen 2011; Walker et al. 2018).

Oat blue dwarf virus

(OBDV)

Taxonomic position Genus: Marafivirus

H

Family: Tymoviridae

OBDV infection in plants of Hordeum vulgare was reported from Czechoslovakia, Finland, North Dakota, and Minnesota (USA), and Sweden (Banttari and Moore 1962; Westdal 1968; Timian 1985). The virus-infected barley plants exhibit symptoms of blue green coloration and stunting, tiller excessively and either do not produce heads or have sterile spikes. Leaves are short and stiff and have enations along the veins of the leaves and leaf sheaths. The virus is transmitted primarily by a adult aster leafhopper, Macrosteles fascifrons, with occasional transmission by immature leafhoppers in a persistent-propagative manner. The virus cannot be transmitted by seed, mechanically, or soil. For more details of OBDV, refer to Avena sativa.

Oat sterile dwarf virus Taxonomic position Genus: Fijivirus

(OSDV)

Family: Reoviridae

OSDV infection in plants of Hordeum vulgare was reported from England and Europe (Catherall 1970). The virus-infected barley plants exhibit symptoms of darker green coloration, slight stunting, and production of few juvenile tillers. The virus is transmitted by delphacid planthoppers, especially Javesella pellucida and Dicranotropis hamata in a propagative manner. For more details of OSDV, refer to Avena sativa.

Rice black streaked dwarf virus Taxonomic position Genus: Fijivirus

(RBSDV)

Family: Reoviridae

RBSDV infection in plants of Hordeum vulgare was reported from China, Japan, Korea, and Russia (Zhou et al. 2010). The virus-infected barley plants exhibit veinal swellings, severe stunting and leaf

1192

Hordeum vulgare (Barley)

twisting. The virus is transmitted by the planthopper vectors, Laodelphax striatellus, Unkanodes sapporonus, and U. albifascia, in a propagative manner but not by seed or sap. For more details of RBSDV, refer to Oryza sativa.

Soil-borne wheat mosaic virus Taxonomic position Genus: Furovirus

(SBWMV)

Family: Virgaviridae

SBWMV infection in plants of Hordeum vulgare was reported from Argentina, Brazil, Canada, Egypt, Italy, Japan, and the USA (Cadle-Davidson and Gray 2006). The virus-infected barley plants exhibit symptoms, which are most prominent in spring on the youngest or lowest leaves, and range from light green to yellow leaf mosaics or mottling that later develops into parallel streaks. Plants may be slightly or severely stunted and rosetted by some strains of SBWMV. Warm weather prevents disease development, thus confining symptoms to lower leaves. The virus is transmitted by the soil-borne fungal vector, Polymyxa graminis, and is also transmissible by mechanical sap-inoculation. The virus is not transmitted by seed. For more details of SBWMV, refer to Triticum aestivum.

Sugarcane yellow leaf virus Taxonomic position Genus: Polerovirus

(ScYLV)

Family: Luteoviridae

ScYLV infection in plants of Hordeum vulgare was reported from Tunisia (Bouallegue et al. 2014). The virus is transmitted by aphid vectors, Melanaphis sacchari and Rhopalosiphum maidis, in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. For more details of ScYLV, refer to Saccharum officinarum.

Wheat dwarf virus Taxonomic position Genus: Mastrevirus

(WDV)

Family: Geminiviridae

WDV infection in plants of Hordeum vulgare was reported from Bulgaria, Czechoslovakia, Russia, China, Spain, Syria, Hungary, and Sweden (Najar et al. 2000; Achon et al. 2006; Koklu et al. 2007; Wang et al. 2008; Tobias et al. 2009, 2010; Ekzayez et al. 2011). Barley plants infected as seedlings are severely stunted and do not form heads, but plants infected later are less stunted. Leaves have small, light green to yellow-brown spots and blotches that may coalesce to cause yellowing and necrosis. The virus is transmitted by the leafhopper vector, Psammotettix alienus, in a persistent, circulative, and non-propagative manner (Ekzayez et al. 2011). The virus is not transmissible by mechanical sapinoculation, and nor transmissible by contact between plants. The virus is not transmitted by seed and pollen. For more details of WDV, refer to Triticum aestivum.

Hordeum vulgare (Barley)

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Wheat streak mosaic virus Taxonomic position Genus: Tritimovirus

(WSMV)

Family: Potyviridae

WSMV infection in plants of Hordeum vulgare was reported from Europe, Syria, Lebanon, North America, Australia, and Russia (Makkouk and Jarikji 1983; Kudela et al. 2008; Maligate et al. 2016). The most severe symptoms occur in barley plants sown early in the autumn but do not appear until the following spring. Initially, light green to light yellow blotches, dashes, or streaks occur parallel to leaf veins, and turn brown as plants mature. Plants become stunted, display a general yellow mottling, and develop large numbers of tillers that vary in height. Plants may die before maturity or have sterile or partially sterile heads with shriveled kernels. At harvest, stunted plants, some with sterile heads, remain that are the same in height or shorter than stubble. The virus is transmitted primarily by wheat curl mites, Aceria tulipae, in a semi-persistent manner, for up to 2.4 km and is also transmissible by mechanical sap-inoculation. For more details of WSMV, refer to Triticum aestivum.

Wheat yellow leaf virus Taxonomic position Genus: Closterovirus

(WYLV)

Family: Closteroviridae

The virus infection in plants of Hordeum vulgare was reported from Italy and Japan (Inouye et al. 1973). The virus-infected barley plants exhibit symptoms of leaves becoming yellow and blistering; plants die or ripen prematurely. The virus is transmitted by the corn leaf aphid, Rhopalosiphum maidis, in a semi-persistent manner. The virus is not transmitted by seed. For more details of WYLV, refer to Triticum aestivum.

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Greber RS (1977) Cereal chlorotic mottle virus (CCMV) a rhabdovirus of Gramineae transmitted by the leafhopper Nesoclutha pallida. Aust Pl Pathol Soc Newsl 6:17 Greber RS (1979) Cereal chlorotic mottle virus – a rhabdovirus of Gramineae in Australia transmitted by Nesoclutha pallida. Aust J Agric Res 30:433–443 Greber RS (1981) Ecological aspects of Cereal chlorotic mottle virus. Aust Pl Pathol Soc Newsl 10:29–30 Greber RS (1982) Cereal chlorotic mottle virus. CMI/AAB Descriptions of Plant Viruses. No. 251 Grylls NE (1963) A striate mosaic virus disease of grasses and cereals in Australia transmitted by the cicadellid Nesoclutha obscura. Aust J Agric Res 14:143–152 Gustafson G, Armour SL (1986) The complete nucleotide sequence of RNA beta from the type strain of barley stripe mosaic virus. Nucleic Acids Res 14(9):3895–3909 Gustafson G, Hunter B, Hanau R, Armour SL, Jackson AO (1987) Nucleotide sequence and genetic organization of barley stripe mosaic virus RNA gamma. Virology 158(2):394–406 Gustafson G, Armour SL, Gamboa GC, Burgett SG, Shepherd JW (1989) Nucleotide sequence of barley stripe mosaic virus RNA alpha: RNA alpha encodes a single polypeptide with homology to corresponding proteins from other viruses. Virology 170(2):370–377 Halbert SE, Pike KS (1986) First report of RMV-type Barley Yellow Dwarf Virus in Barley in Eastern Washington. Plant Dis 70:475 Hamilton RI (1965) An embryo test for detecting seed-borne Barley stripe mosaic in barley. Phytopathology 55:798–799 Hammond J, Lister RM, Foster JE (1983) Purification, identity and some properties of an isolate of Barley yellow dwarf virus from Indiana. J Gen Virol 64:667–676 Harder DE (1975) Electron microscopy of African cereal streak diseased plants. Can J Bot 53:565–581 Harder DE, Bakker W (1973) African cereal streak, a new disease of cereals in East Africa. Phytopathology 63:1407–1411 Hariri D, Fouchard M, Lapierre H (1990) Resistance to barley yellow mosaic virus and to barley mild mosaic virus in barley. In: Koenig R (ed) Proceedings of the first symposium of the international working group on plant viruses with fungal vectors. Verlag Eugen Ulmer, Stuttgart, pp 109–112 Herrera GM, Beratto EM, Andrade OV, Madariaga MV (2001) Identification of Barley stripe mosaic virus (BSMV) on barley in Chile. Agricultura Tecnica 61(3):275–280 Hill AS (1980) Barley yellow mosaic in England. In: 3rd conference on virus diseases of Gramineae in Europe, Rothamsted, 28–30 May Hill SA, Evans EJ (1980) Barley yellow mosaic virus. Plant Pathol 29(4):197–198 Huth W (1990) The yellow mosaic inducing viruses of barley in Germany. In: Proceedings of the 1st symposium of the international working group on plant viruses with fungal vectors, Braunschweig, pp 113–115 Huth W, Lesemann DE, Paul HL (1984) Barley yellow mosaic virus: purification, electron microscopy, serology and other properties of two types of the virus. Phytopathol Z 111:37–54 Huth W, Lesemann DE, Gotz R, Vetten HJ, Mai E, Proeseler G, Signoret P (1995) Brome streak mosaic virus isolated from Barley in South France. Agron EDP Sci 15:510 Hyung LS, Shikata E (1977) Occurrence of Northern cereal mosaic virus in Korea. Plant Prot 16:87–92 Inouye T, Mitsuhata K, Heta H, Hiura U (1973) A new virus of wheat, barley and several other plants in Gramineae, Wheat yellow leaf virus. Nogaku Kenkyu Ohara Inst Agric Biol, Okayama Univ 55:1–6 Ito S, Fukushi T (1944) Studies on northern cereal mosaic. J Sapporo Soc Agric For 36:62–89 Jackson AO, Lane LC (1981) Hordeiviruses. In: Hand Book of Plant Virus Infections: Comparative Diagnosis. Kurstak E (ed). Elsevier/North Holland Biomedical Press, Oxford, pp 565–625 Jackson AO, Bragg J, Lim HS, Ganesan U (2011) Hordeivirus. In: Tidona C, Darai G (eds) The Springer index of viruses. Springer, New York. https://doi.org/10.1007/978-0-387-95919-1 Jedlinski H (1981) Rice root aphid Rhopalosiphum rufiabdominalis, a vector of Barley yellow dwarf virus in Illinois, and the disease complex. Plant Dis 65:975–978 Jezewska M (2001) Identification of Barley stripe mosaic virus in Poland. J Plant Prot Res 41:164–167 Jezewska M, Trzmiel K (2009) First report of Barley yellow mosaic virus infecting barley in Poland. Plant Pathol 58:784 Jezewska M, Trzmiel K (2010) Studies on cereal soil-borne viruses in Poland. J Plant Protect Res 50:527–534 Jianping C, Swaby AG, Adams MJ, Yili R (1991) Barley mild mosaic virus inside its fungal vector, Polymyxa graminis. Ann Appl Biol 118:615–621 Kashiwazaki S (1996) The complete nucleotide sequence and genome organization of Barley mild mosaic virus (Na1 strain). Arch Virol 141:2077–2089 Kashiwazaki S, Ogawa K, Usugi T, Omura T, Tsuchizaki T (1989) Characterization of several strains of Barley yellow mosaic virus. Ann Phytopathol Soc Jpn 55:16–25 Kashiwazaki S, Minobe Y, Omura T, Hibino H (1990) Nucleotide sequence of Barley yellow mosaic virus RNA 1: a close evolutionary relationship with potyviruses. J Gen Virol 71:2781–2790

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1196

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Kashiwazaki S, Minobe Y, Hibino H (1991) Nucleotide sequence of Barley yellow mosaic virus RNA 2. J Gen Virol 72:995–999 Katis N, Tzavella-Klonari K, Adams MJ (1997) Occurrence of barley yellow mosaic and barley mild mosaic bymoviruses in Greece. Eur J Plant Pathol 103:281–284 Koklu G, Ramsell JNE, Kvarnheden A (2007) The complete genome sequence for a Turkish isolate of Wheat dwarf virus (WDV) from barley confirms the presence of two distinct WDV strains. Virus Genes 34:359–366 Kudela O, Kudelova M, Novakova S, Glasa M (2008) First report of Wheat streak mosaic virus in Slovakia. Plant Dis 92:1365 Kumari SG, Muharram I, Makkouk KM, Al-Ansi A, El-Pasha R, Al-Motwkel WA, Haj Kassem A (2006) Identification of viral diseases affecting barley and bread wheat crops in Yemen. Aust Plant Pathol 35:563–568 Kundu JK (2008) First report of Barley yellow dwarf virus-PAS in Wheat and Barley Grown in the Czech Republic. Plant Dis 92:1587 Kundu JK (2009) First report of Barley yellow dwarf virus-MAV in Oat, Wheat, and Barley Grown in the Czech Republic. Plant Dis 93:964 La YJ, Park YK (1979) Serological detection of Barley stripe mosaic virus infection in the seeds of barley and wheat cultivars grown in Korea. Korean J Plant Pathol 18:29–33 Lee K, Kashiwazaki S, Habi T, So I (1996) Properties and capsid protein gene sequence of a Korean isolate of Barley mild mosaic virus. Ann Phytopath Soc Japan 62:397–401 Lister RM, Ranieri R (1995) Distribution and economic importance of Barley yellow dwarf. In: D’Arcy CJ, Burnett PA (eds) Barley yellow dwarf: 40 years of progress. American Phytopathological Society, St. Paul, pp 29–53 Lister RM, Carroll TW, Zaske SK (1981) Sensitive serologic detection of Barley stripe mosaic virus in barley seed. Plant Dis 65:809–814 Lockhart BEL (1986) Occurrence of Cereal chlorotic mottle virus in northern Africa. Plant Dis 70:912–915 Lockhart BEL, El Maataoui M, Carroll TW, Lennon AM, Zaske SK (1986) Identification of Barley yellow striate mosaic virus in Morocco and its field detection by enzyme immune assay. Plant Dis 70:1113–1117 Lopez-Moya JJ, Valli A, Garcia JA (2009) Potyviridae. In: Encyclopaedia of Life Sciences (ELS). Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000755.pub2 Makkouk KM (2003) Transmission of Barley yellow striate mosaic virus (BYSMV) by the planthopper vector Laodelphax striatella (Fallen) in Syria and Lebanon. Eighth Arab Congress of Plant Protection, El-Beida, 12–16 Oct 2003 Makkouk KM, Jarikji DA (1983) Detection of sap-transmissible viruses infecting cereals in Jordan, Lebanon and Syria (Brome mosaic virus, Barley stripe mosaic virus, Wheat streak mosaic virus). Z Pflanzenkrankh Pflanzenschuts J. Plant Dis Prot 90:12–17 Makkouk KM, Bertschinger L, Conti M, Bolay N, Dusunceli F (1996) Barley yellow striate mosaic rhabdovirus naturally infects cereal crops in the Anatolian Plateau of Turkey. J Phytopathol 144:413–415 Makkouk KM, Najar A, Kumari SG (2001a) First record of Barley yellow dwarf and Cereal yellow dwarf viruses in Tunisia. Plant Pathol 50(6):806 Makkouk KM, Ghulam W, Kumari SG (2001b) First report of Barley yellow striate mosaic virus infecting Barley and Wheat in Lebanon. Plant Dis 85:441 Maligate A, Adorada D, Chambers G, Terras MA (2016) Occurrence of winter cereal viruses in New South Wales, Australia, 2006 to 2014. Plant Dis 100:313–317 Malmstrom CM, Shu R (2004) Multiplexed RT-PCR for streamlined detection and separation of barley and cereal yellow dwarf viruses. J Virol Methods 120(1):69–78 Martin RR, D’Arcy CJ (1995) Taxonomy of Barley yellow dwarf viruses. In: D’Arcy CJ, Burnett PA (eds) Barley yellow dwarf: 40 years of progress. American Phytopathological Society, St. Paul, pp 203–214 Miller MA, Rosachova L (1997) Barley yellow dwarf viruses. Annu Rev Phytopathol 35:167–190 Mirshekar A, Rasoulian GR, Mosahebi GH (2013) Efficiency of ten populations of Schizaphis graminum (Rondani) in the transmission of barley yellow dwarf virus-PAV (BYDV-PAV) in Iran. African J Plant Sci 7:118–123 Nagaich BB, Vashisth KS (1963) Barley yellow dwarf: a new viral disease for India. Indian Phytopath 16:318–319 Najar A, Makkouk KM, Kumari SG (2000) First record of Barley yellow striate mosaic virus, Barley stripe mosaic virus, and Wheat dwarf virus infecting cereal crops in Tunisia. Plant Dis 84:1045 Najar A, Hamdi I, Varsani A (2017) Barley yellow dwarf virus in barley crops in Tunisia: prevalence and molecular characterization. Phytopathol Mediterr 56:111–118 Nomura K, Kashiwazaki S, Hibino H, Inoue T, Nakata E, Tsuzaki Y, Okuyama S (1996) Biological and serological properties of two strains of Barley mild mosaic virus. J Phytopathol 144:103–107 Ogawa, Moichi (1984) Ann Phytopath Soc Japan 50:108 Oswald JW, Houston BR (1951) A new virus disease of cereals, transmissible by aphids. Plant Dis Reptr 35:471–475 Panayotou PC (1980) Occurrence of barley yellow dwarf virus in Greece. Phytopathol Mediterr 19:143–144

Hordeum vulgare (Barley)

1197

Peerenboom E, Prols M, Schell J, Steinbiss HH, Davidson AD (1992) The complete nucleotide sequence of RNA 1 of a German isolate of Barley yellow mosaic virus and its comparison with a Japanese isolate. J Gen Virol 73:1303–1313 Peerenboom E, Jacobi V, Antoniw JF, Schlichter UHA, Cartwright EJ, Steinbiss HH, Adams MJ (1996) The complete nucleotide sequence of RNA-2 of a fungally – transmitted UK isolate of Barley mild mosaic bymovirus (BaMMV) and identification of amino acid combinations possibly involved in fungus transmission. Virus Res 40:149–159 Pocsai E (1987) Effect of Brome mosaic virus infection on the plant height and weight of cereals at their early stages of growth. Cereals Res Com 15:167–174 Pocsai E, Kobza S, Muranyi I, Szunics L (1991) Brome mosaic virus infection in different cereal breeding materials. Acta Phytopathol Entomol Hung 26:207–212 Power AG, Gray SM (1995) Aphid transmission of Barley yellow dwarf viruses: interactions between viruses, vectors, and host plants. Barley yellow dwarf-40 years of progress, D’Arcy C, Burnett P., Am Phytopath Soc Press, St-Paul, 259–289. Rabenstein F, Huss H (2013) Studies on grass viruses in Austria. In: 63. Jahrestagung der Vereinigung der Pflanzenzüchter und Saatgutkaufleute Österreichs, Raumberg-Gumpenstein, pp 15–18 Ramel ME, Brown DJF, Valloton R, McGavin W, Gugerli P (1994) Occurrence of infection by a strain of Arabis mosaic virus nepovirus of barley (cv. Express) and viruliferous Xiphinema diversicaudatum in fields near Fribourg in Switzerland. Proc 22nd International Nematology Symposium. 7–12 August 1994. University Gent Zoology Institute, KL Ledeganckstraat 35, B-9000 Gent 106–107 Robertson NL (2003) Luteovirus and polerovirus found in small grains for the first time in the Matanuska-Susitna Region of Alaska. Plant Dis 87:446 Robertson NL, Brumfield SK (2000) First report of Barley yellow streak mosaic virus infected barley in Alaska. Plant Dis 84:595 Robertson NL, Carroll TW (1988) Virus-like particles and a spider mite intimately associated with a new disease of barley. Science 240:1188–1190 Robertson NL, Carroll TW (1989) Electron microscopy of the novel Barley yellow streak mosaic virus. J Ultrastruct Mol Struct Res 102:139–146 Robertson NL, Carroll TW (1991) Mechanical transmission, partial purification and preliminary chemical analysis of Barley yellow streak mosaic virus. Plant Dis 75:839–843 Rochow WF (1969) Biological properties of four isolates of barley yellow dwarf virus. Phytopathology 59:1580–1589 Rochow WF (1979) Field variants of Barley yellow dwarf virus: detection and fluctuation during twenty years. Phytopathology 69:655–660 Ruan YL, Lin DD, Xu RY (1983) Transmission characteristics of the Northern cereal mosaic virus by the small brown planthopper (Laodelphax striatellus Fallen). Acta Phytopathol Sin 13:20–24 Salm SN, Rey MEC, Robertson NL, French R, Rabenstein F, Schubert J (1996) Molecular cloning and nucleotide sequencing of the partial genomes of Agropyron and Hordeum mosaic viruses, two members of the Rymovirus genus in the taxonomic family Potyviridae. Arch Virol 141:2115–2127 Shepherd DN, Martin DP, Van Der Walt E, Dent K, Varsani A, Rybicki EP (2010) Maize streak virus: an old and complex ‘emerging’ pathogen. Mol Plant Pathol 11(1):1–12 Shikata E, Lu YT (1967) Electron microscopy of Northern cereal mosaic virus in Japan. Proc Jpn Acad 43:918–923 Skaf JS, Carroll TW (1995) Purification of Barley yellow streak mosaic virus and detection by DAS-ELISA and ISEM using polyclonal antibodies. Plant Dis 79:1003–1007 Skaf JS, Brumfield SK, Carroll TW, Forster RL (1992) Barley yellow streak mosaic virus infection of barley in Idaho. Plant Dis 76:861 Smidansky ED, Carroll TW (1996) Factors influencing the outcome of Barley yellow streak mosaic virus brown wheat mite – barley interactions. Plant Dis 80:186–193 Smith O, Clapham A, Rose P, Liu Y, Wang J, Allaby RG (2014) A complete ancient RNA genome: identification, reconstruction and evolutionary history of archaeological Barley stripe mosaic virus. Sci Rep 4:4003. https://doi.org/ 10.1038/srep04003 Svanella-Dumas L, Candresse T, Hulle M, Marais A (2013) Distribution of Barley yellow dwarf virus-PAV in the SubAntarctic Kerguelen Islands and characterization of two new luteovirus species. PLoS One 8(6):E67231 Tanno F, Nakatsu A, Toriyama S, Kojima M (2000) Complete nucleotide sequence of Northern cereal mosaic virus and its genome organization. Arch Virol 145:1373–1384 Timian RG (1985) Oat blue dwarf virus in its plant host and insect vectors. Plant Dis 69:706–708 Timpe U, Kuhne T (1994) The complete sequence of RNA 2 of Barley mild mosaic virus (BaMMV). Eur J Plant Pathol 100:233–241 Tobias I, Kiss B, Bakardjieva N, Palkovics L (2009) The nucleotide sequence of barley strain of Wheat dwarf virus isolated in Bulgaria. Cereal Res Commun 37(2):237–242 Tobias I, Kiss B, Salanki K, Palkovics L (2010) The nucleotide sequence of barley strain of Wheat dwarf virus isolated in Hungary. Cereal Res Commun 38(1):67–74

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Hosta spp.

Trzmiel K (2017) Identification of Barley yellow dwarf viruses in Poland. J Plant Pathol 99(2). https://doi.org/10.4454/ jpp.v99i2.3869 Ueng PP, Vincent JR, Kawata EE, Lei CH, Lister RM, Larkins BA (1992) Nucleotide sequence analysis of the genomes of the MAV-PS1 and P-PAV isolates of barley yellow dwarf virus. J Gen Virol 73:487–492 Vincent JR, Ueng PP, Lister RM, Larkins BA (1990) Nucleotide sequences of coat protein genes for three isolates of barley yellow dwarf virus and their relationships to other luteovirus coat protein sequences. J Gen Virol 71(12):2791–2799 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Rhabdoviridae. J Gen Virol 99:447–448 Wang XF, Wu B, Wang JF (2008) First report of Wheat dwarf virus infecting barley in Yunnan, China. J Plant Pathol 90:400 Westdal PH (1968) Host range studies of Oat blue dwarf virus. Can J Bot 46:1431–1435 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Yan T, Zhu JR, Di D, Gao Q, Zhang Y, Zhang A, Yan C, Mial H, Wang XB (2015) Characterization of the complete genome of Barley yellow striate mosaic virus reveals a nested gene encoding a small hydrophobic protein. Virology 478:112–122 Zagula KR, Niblett CL, Robertson NL, French R, Lommen SA (1992) Potyviridae: genus Rymovirus. Arch Virol 5:269–276 Zamora MR, Burnett PA, Vivar HE, Rodriguez R, Navarro M (1988) Barley stripe mosaic virus in Mexico. Plant Dis 72:546 Zhang QF, Guan WN, Ren ZY, Zhu XS, Tsai JH (1983) Transmission of Barley yellow dwarf virus strains from northwestern China by four aphid species. Plant Dis 67:895–899 Zhang W, Cheng Z, Xu L, Wu M, Waterhouse P, Zhou G, Li S (2009) The complete nucleotide sequence of the barley yellow dwarf GPV isolate from China shows that it is a new member of the genus Polerovirus. Arch Virol 154:1125–1128 Zheng T, Cheng Y, Chen JP, Antoniw JF, Adams MJ (1999) The occurrence of Barley mild mosaic virus (BaMMV) in China and the nucleotide sequence of its coat protein gene. J Phytopathol 147:229–234 Zhou T, Wang Y, Fan YJ, Cheng ZB, Zhou YJ (2010) First report of Rice black-streaked dwarf virus infecting barley in Jiangsu, China. J Plant Pathol 92:S4.118

Hosta spp. Family: Asparagaceae

Ornamental

Arabis mosaic virus

(ArMV)

Taxonomic position Genus: Nepovirus

Family: Secoviridae

ArMV infection in plants of Hosta spp. was reported from North America (Lockhart 2006). The virusinfected hosta plants exhibit symptoms of leaf deformation and stunting. The virus is transmitted by nematode vectors (Xiphinema spp.) in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

Hosta spp.

1199

CMV infection in plants of Hosta spp. was reported from Ohio (USA) (Fisher 2013b). The virusinfected hosta plants exhibit mottle and spotting symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Hosta virus X Taxonomic position Genus: Potexvirus

(HVX)

Family: Alphaflexiviridae

Geographical distribution HVX infection in plants of Hosta spp. was first identified in 1996 in Minnesota, Indiana, Iowa, New Zealand, and Michigan (USA) (Currier and Lockhart 1996; Baker 2013). Since this report, it has been found all around the world in countries such as Germany, Japan, Korea, New Zealand, the USA, the Czech Republic, the UK, and China (Blanchette and Lockhart 2003; Ryu et al. 2006; Adedire et al. 2009; Kolackova et al. 2011; Richert-Poeggeler et al. 2012; Tang et al. 2012; Wei et al. 2013). Symptoms and host(s) The virus-infected hosta plants exhibit symptoms ranging from latency to severe mosaic and leaf necrosis. The symptoms also included leaf thickening and blotching and abnormal variegation and various patterns of leaf discolorization. In some cultivars mottling and interveinal chlorosis between secondary veins are noticed. Leaf desiccation and necrotic leaf tissue may also be observed. Flower breaking symptoms were also noticed in flowers of infected plants (Currier and Lockhart 1996). Wei et al. (2013) have observed hosta plants showing leaf deformation, puckering, and ink-bleed symptoms. Some varieties do not show symptoms, and newly infected plants may not show typical symptoms for months or even years after the initial infection. Transmission The virus is mechanically sap-transmissible. As far as is known, Hosta spp. are the only natural hosts for this virus, and all Hosta spp. appear to be susceptible. The virus also spreads by using virus-infected vegetative propagative material. The virus spreads from plant to plant when the sap of an infected plant comes into contact with a wound on a healthy plant. These wounds are made with tools used in propagation and upkeep, such as pruning shears, string trimmers, and lawn mowers (Kennelly 2007). It can also be moved by human hands when plants are divided or groomed (e.g., removing old leaves or flowers). The virus was seed-transmitted up to 7.5% in hosta (Ryu et al. 2006). No vector is reported for this virus. Nicotiana benthamiana could be infected experimentally by transcripts from an infectious cDNA clone of HVX (De La Torre et al. 2012). Virion properties and genome The virions are flexuous filaments, 530 nm in length, and 13 nm in diameter (Richert-Poeggeler et al. 2012). The genome consists of a single linear molecule of positive-sense ssRNA of 6528 nt (AJ620114 = NC_011544) and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type, 18–27 kDa in size (Park and Ryu 2003; Adams et al. 2004).

H

1200

Hosta spp.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Hosta spp. was reported from the USA (Lockhart and Currier 1996). The virus-infected hosta plants show primary foliar lesions followed by latent infection. The virus is transmitted by a thrips vector, Frankliniella occidentalis, in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tobacco rattle virus

(TRV)

Taxonomic position Genus: Tobravirus

Family: Virgaviridae

TRV infection in plants of Hosta spp. was reported from Europe, the USA, and the UK (Lockhart et al. 1995; Lockhart and Currier 1996; Harju et al. 2011; Koenig et al. 2012; Fisher 2013c). The virusinfected hosta plants exhibit leaf mottle symptoms. The virus is transmitted by nematode vectors and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Hosta spp. was reported from the USA and Lithuania (Lockhart and Currier 1996). The virus-infected hosta plants exhibit chlorosis symptoms on the leaves. The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Hosta spp. was reported from the USA (Fisher 2013a). The virus-infected hosta plants exhibit mottle symptoms. The virus is transmitted by the thrips vectors; the virus present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Hosta spp.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

1201

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Hosta spp. was reported from the USA (Lockhart and Currier 1996; Navalinskiene and Samuitiene 2000; Samuitien_e and Navalinskien_e 2001). The virus-infected hosta plants produce symptoms of chlorotic spotting, leaf mottling, and chlorosis. The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ToRSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV has been reported to infect plants of Hosta spp. in the USA, causing leaf necrosis and stunting, chlorotic and necrotic spotting, and distinctive ringspots (Lockhart and Currier 1996; Momol et al. 2003). The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Adedire OL, Wen R-H, Windham A, Windham M, Hajimorad MR (2009) Hosta virus X in Hosta identified in Tennessee, USA. Plant Pathol 58:405 Baker CA (2013) Hosta virus X, a Potexvirus. Plant pathology circular no. 410. Florida Deparment of Agriculture and Consumer Services, Division of Plant Industry Blanchette B, Lockhart B (2003) Hosta virus X: a three-year study. Hosta J 35:19–23 Currier S, Lockhart BEL (1996) Characterization of a potexvirus infecting Hosta spp. Plant Dis 80:1040–1043 De La Torre CM, Qu F, Redinbaugh MG, Lewandowski DJ (2012) Biological and molecular characterization of a U.S. isolate of Hosta virus X. Phytopathology 102:1176–1181 Fisher JR (2013a) Identification of Tobacco streak virus associated with a virus-like mottle symptom on Hosta. Plant Health Prog. https://doi.org/10.1094/PHP-2013-122-01-BR Fisher JR (2013b) Identification of a Cucumber mosaic virus subgroup II strain associated with virus-like symptoms on Hosta in Ohio. Online. Plant Health Prog. https://doi.org/10.1094/PHP-2013-123-01-BR Fisher JR (2013c) Identification of two Tobacco rattle virus sequence variants associated with virus-like mottle symptom on Hosta in Ohio. Online. Plant Health Prog. https://doi.org/10.1094/PHP-2013-0330-01-RS Harju VA, Skelton AL, Monger WA, Forde SMD, Daly M, Nixon T, Lawson R, Bennett S, Mumford RA (2011) A report of viruses, viroids and phytoplasmas found in ornamental plants from 1999–2007: findings from central science laboratory, UK. Acta Hortic 901:223–229 Kennelly MM (2007) Hosta virus X.EP-192, Kansas State University Agricultural Experiment Station and Cooperative Extension Service. www.plantpath.k-state.edu/doc/extension-factsheets/hosta-virus-X.pdf Koenig R, Lesemann DE, Pleij CWA (2012) Tobacco rattle virus genome alterations in the Hosta hybrid ‘Green Fountain’ and other plants: reassortments, recombinations and deletions. Arch Virol 157:2005–2008 Kolackova A, Safrankova I, Holkova L (2011) First report of Hosta virus X infecting Hosta in the Czech Republic. New Dis Rep 24:9 Lockhart BEL (2006) Occurrence of Arabis mosaic virus in Hostas in the United States. Plant Dis 90:834 Lockhart BEL, Currier S (1996) Viruses occurring in Hosta spp. in the USA. Acta Hortic 432:62–71

H

1202

Hovea spp. (Hovea elliptica and H. pungens)

Lockhart BE, Morelli G, Westendorp J (1995) Occurrence of tobravirus infections in Peperomia, Hosta, and Phlox in the midwestern U.S. Plant Dis 79:1249 Momol MT, Dankers H, Adkins S (2003) First report of tomato spotted wilt virus in hosta in Florida. Online. Plant Health Prog. https://doi.org/10.1094/PHP-2003-1024-01-HN Navalinskiene M, Samuitiene M (2000) Natural occurrence of tomato ringspot nepovirus in ornamental plants in Lithuania. In: Proceedings of the international conference: development of environmentally friendly plant protection in the Baltic Region, Tartu, pp 140–143, 28–29 Sept 2000 Park MH, Ryu KH (2003) Molecular evidence supporting the classification of Hosta virus X as a distinct species of the genus. Arch Virol 148(10):2039–2045 Richert-Poeggeler K, Commandeur U, Kobayashi K, Maass C, Lockhart B (2012) Electron microscopic studies of vegetative and generative tissues from Hosta virus X infected hosta. Bioforsk Fokus 7(9):37 Ryu KH, Park MH, Lee MY, Lee JS (2006) Characterization and seed transmission of Hosta virus X isolated from Hosta plants. Acta Hortic 722:91–93 Samuitien_e M, Navalinskien_e M (2001) Nepoviruses and their influence on field floriculture. Biologija 4:43–45 Tang J, Hardy C, Lebas BSM, Ward LI (2012) Presence of Hosta virus X in New Zealand. Australasian Plant Dis Notes 7: 39–40 Wei MS, Zhang YJ, Li GF, Ma J, Li M (2013) First report of Hosta virus X infecting hosta plants in China. Plant Dis 97(3):429

Hovea spp. (Hovea elliptica and H. pungens) Family: Fabaceae

Ornamental

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Hovea spp. was reported from Australia (McKirdy et al. 1994; Webster et al. 2007). The virus-infected hovea plants exhibit mosaic and stunting symptoms. The virus is transmitted by aphid vectors in a non-persistent manner and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

References McKirdy SJ, Coutts BA, Jones RAC (1994) Occurrence of Bean yellow mosaic virus in subterranean clover pastures and perennial native legumes. Aust J Agric Res 45:183–194 Webster CG, Coutts BA, Jones RAC, Jones MGK, Wylie SJ (2007) Virus impact at the interface of an ancient ecosystem and a recent agroecosystem: studies on three legume-infecting potyviruses in the southwest Australian floristic region. Plant Pathol 56:729–742

Hoya spp. (Waxflower) Family: Apocynaceae

Ornamental

Capsicum chlorosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(CaCV)

Family: Tospoviridae

Hoya spp. (Waxflower)

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CaCV was reported in plants of Hoya calycina in Brisbane (Australia) and the USA (Persley et al. 2006; Melzer et al. 2014). The virus-infected waxflower plants exhibit ringspots, line patterns, chlorotic blotches, and necrotic etching symptoms. The virus is transmitted by thrips vectors in a persistentpropagative manner, and is also transmissible by mechanical sap-inoculation. For more details of CaCV, refer to Capsicum annuum.

Hoya chlorotic spot virus

(HCSV)

Taxonomic position HCSV is a tentative member of the genus Tobamovirus and family Virgaviridae Geographical distribution HCSV infection in plants of Hoya wayetii was reported from the USA (KX434725; Adkins et al. 2018). Transmission The virus has no known natural vector. The virus is mechanically sap-transmissible. Virion properties and genome Virions are 18 nm in diameter and have a predominant length of 300 nm. The genome consists of a positive-sense single-stranded RNA of 6386 nt (KX434725) (Adams et al. 2017; Adkins et al. 2018).

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV was reported infecting plants of Hoya australis from Germany (Koenig 1985). There is no known vector for this virus. The virus is mechanically sap-transmissible and also transmissible by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tomato chlorotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(TCSV)

Family: Tospoviridae

TCSV has been detected in plants of Hoya wayetii in Florida (Baker and Adkins 2015). The virusinfected waxflower plants exhibit necrotic spots, rings, and line pattern symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation. For more details of TCSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

H

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Humulus japonicus (Japanese hop)

TSWV infection in plants of Hoya carnosa was reported from Pennsylvania (USA) (Hausbeck et al. 1992). The virus-infected waxflower plants exhibit necrosis and chlorotic ringspot symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Adams MJ, Adkins S, Bragard C, Gilmer D, Li D, MacFarlane SA, Wong S-M, Melcher U, Ratti C, Ryu KH, and ICTV Report Consortium (2017) ICTV virus taxonomy profile: Virgaviridae. J Gen Virol 98:1999–2000 Adkins S, D’Elia T, Fillmer K, Pongam P, Baker CA (2018) Biological and genomic characterization of a novel Tobamovirus infecting Hoya spp. Plant Dis 102(12):2571–2577 Baker CA, Adkins S (2015) First report of Tomato chlorotic spot virus in Hoya wayetii and Schlumbergera truncata. Plant Health Prog. https://doi.org/10.1094/PHP-BR-14-0043 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among greenhouse ornamentals in Pennsylvania. Plant Dis 76:795–800 Koenig R (1985) Recently discovered virus or virus-like diseases of ornamentals and their epidemiological significance. Acta Hortic 164:21–31 Melzer M, Shimabukuro J, Long M, Nelson SC, Alvarez AM, Borth WB, Hu J (2014) First report of Capsicum chlorosis virus infecting waxflower (Hoya calycina Schlecter) in the United States of America. Plant Dis 98:571 Persley DM, Thomas JE, Sharman M (2006) Tospoviruses an Australian perspective. Australas Plant Pathol 35:161–180

Humulus japonicus (Japanese hop) Synonyms Humulus scandens Family: Cannabaceae

Ornamental

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

BWYV infection in plants of Humulus scandens was reported from China (Wang et al. 2013). The virus-infected Japanese hop plants show mosaic symptoms of young leaves. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of BWYV, refer to Beta vulgaris.

Humulus japonicus latent virus Synonyms Humulus japonicus virus

(HJLV)

Humulus lupulus (Hop)

Taxonomic position Genus: Ilarvirus

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Family: Bromoviridae

Geographical distribution HJLV infection in plants of Humulus japonicus was reported from the UK and China (Adams et al. 1989). Symptoms and host(s) The virus-infected Japanese hop plants exhibit symptomless infections. Transmission The virus is transmissible by mechanical inoculation to more than nine plant families which show necrotic local lesions and chlorotic mottle or mosaic in infected leaves. The virus is transmitted by seed.

H Virion properties and genome The virions are isometric, non-enveloped, and 24–33 nm in diameter. The genome consists of three segments of positive-sense single-stranded RNAs. The reference sequences of RNA 1 (3382 nt; AY500236 = NC_006064), RNA 2 (2810 nt; AY500237 = NC_006065), and RNA 3 (1938 nt; AY500238 = NC_006066) are available (Scott and Zimmerman 2006; Scott 2011a, b).

References Adams AN, Clark MF, Barbara DJ (1989) Host range, purification and some properties of a new Ilarvirus from Humulus japonicus. Ann Appl Biol 114:497–508 Scott SW (2011a) Bromoviridae and allies. In: Encyclopedia of Life Sciences (ELS). Wiley, Chichester. https://doi.org/ 10.1002/9780470015902 Scott SW (2011b) Ilarvirus. Bromoviridae. In: The Springer Index of Viruses. Springer, New York, pp 187–194. https:// doi.org/10.1007/978-0-387-95919-1_27 Scott SW, Zimmerman MT (2006) The complete sequence of the genome of Humulus japonicus latent virus. Arch Virol 151:1683–1687 Wang R, Wang N, Ye T, Chen SY, Fan ZF, Zhou T (2013) Occurrence of Beet western yellows virus in Japanese hop (Humulus scandens) in China. J Plant Pathol 95:450

Humulus lupulus (Hop) Family: Cannabaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Ornamental

(AMV)

Family: Bromoviridae

AMV infection in plants of Humulus lupulus was reported from China and the former Czechoslovakia (Novak and Lanzova 1976; Yu and Liu 1987; Pethybridge et al. 2008). The virus-infected hop cv. Styrian exhibits systemic chlorotic spotting. The virus is transmitted by a large number of aphid

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Humulus lupulus (Hop)

species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

American hop latent virus Taxonomic position Genus: Carlavirus

(AHLV)

Family: Betaflexiviridae

Geographical distribution AHLV infection in plants of Humulus lupulus was first reported by Probasco and Skotland (1976a). The virus is reported from the UK, Germany, Australia, New Zealand, and the USA (Adams and Barbara 1982; Adams et al. 1982; Munro 1987; Hay et al. 1992; Klein and Husfloen 1995; Pethybridge et al. 2008, 2009; Eastwell and Druffel 2012). Symptoms and host(s) Some British hop cultivars do not display symptoms, whereas American hop cultivars show ring and line patterns in leaves. Hop is the only known natural host of AHLV (Adams et al. 1982). Transmission The virus is transmitted by aphid vectors, Phorodon humuli, Macrosiphum euphorbiae, and Myzus persicae, in a non-persistent manner (Paine 1953; Adams and Barbara 1982; Eppler 1994). The virus is transmissible by mechanical sap-inoculation; Chenopodium quinoa expresses chlorotic spots and systemic vein-banding; and Datura stramonium expresses local lesion reactions suitable for virus diagnosis. Vegetative propagation of virus-infected plants is a major route of virus dissemination. There are no reports of seed transmission of AHLV. Virion properties and genome The virions are flexuous filaments about 680 nm in length and 15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8601 nt (NC_017859) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Probasco and Skotland 1976b; Adams and Barbara 1982; Adams et al. 2004; Eastwell and Druffel 2012).

Apple fruit crinkle viroid

(AFCVd)

Taxonomic position AFCVd is a tentative member of the genus Apscaviroid and family Pospiviroidae AFCVd isolate in cultivated plants of Humulus lupulus was reported from Japan, China, and Washington (Ito et al. 1993; Sano et al. 2004; Pethybridge et al. 2008; Lu et al. 2012). AFCVd shows the roughened and crinkled appearance of apple fruit on cultivars like “Ohrin,” “Mutsu,” “Jonathan,” “Fuji,” etc. and blister bark symptoms of twigs and branches on some cultivars like “Nero 26” and a crab apple “NY58-22” (Ito and Yoshida 1998). AFCVd shows stunting, leaf curling, and reduction of alpha-acid contents in the infected cones of cultivated hops (Sano et al. 2004, 2008). The viroid is sap-transmissible. Once the pathogen is introduced into a hop yard, it is readily spread

Humulus lupulus (Hop)

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mechanically by hands and cutting tools and by contact with vines through horticultural operations such as crowning, stripping, and picking. For more details of AFCVd, refer to Malus domestica.

Apple mosaic virus

(ApMV)

Taxonomic position Genus: Ilarvirus

Family: Bromoviridae

ApMV infection in plants of Humulus lupulus was reported from Australia, Germany, New Zealand, the UK, and the USA (Albrechtova et al. 1979; Sano et al. 1985; Kanno et al. 1993; Klein and Husfloen 1995; Pethybridge et al. 2002a, b, 2008, 2009). Two serotypes of ApMV are found infecting hops, ApMV-H (hop) and ApMV-I (intermediate) (Crowle et al. 2003). The virus-infected hop plants show leaf symptoms of, chlorotic ringspots which later become necrotic, and oak-leaf-like patterns. ApMV in hop can cause significant losses in cone yield and decreased levels of brewing organic acids. Virus infections overall may cause problems in plant propagation, including death of softwood cuttings in hops. No insect vector is known for this virus, although other ilarviruses are transmitted by the interaction between pollen and thrips. The virus is mechanically sap-transmissible, by plant-to-plant contact, pruning, and other contacts during agronomic practices. For more details of ApMV, refer to Malus domestica.

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

A Humulus lupulus strain of ArMV (ArMV-H) has been reported (Adams et al. 1986, 1987) which has been in commercial hop yards in several countries, including Belgium, the former Czechoslovakia, France, Australia, Germany, New Zealand, South Africa, the USA, and the UK (Keyworth and Davies 1946; Cartledge 1956; Bock 1966; Davies and Clark 1983; Thresh and Edwards 1983; Polak and Svoboda 1988; Thresh and Ormerod 1989; Hay et al. 1992; Kominek et al. 2003; Pethybridge et al. 2008, 2009). ArMV-H infection of hop can cause a variety of symptoms including bare bine or spidery hop, split leaf blotch, hop chlorotic disease, and nettlehead (Keyworth and Davies 1946; Thresh and Pitcher 1978; Adams et al. 1987). Symptoms of bare bine or spidery hop are apparent in early spring as weak shoot growth, curvature of shoots, and production of small, dark-colored leaves. Symptoms of split leaf blotch include translucent, yellow oily blotches between the leaf veins, leading to leaves splitting upon expansion (Thresh et al. 1972). Plants with nettlehead often exhibit symptoms of bare bine early in the season. Later, plants develop stiff, erect shoots with short internodes that fail to climb, and which fall away from support strings. Other symptoms of nettlehead include upward rolling of the leaf margins, vein-clearing or mottle, and enations on the abaxial leaf surface (Davies and Clark 1983). Plants with hop chlorotic disease often exhibit symptoms including severe distortion of developing foliage on the lower parts of bines, with leaves often assuming a characteristic parrot beak shape. ArMV-H is transmitted by the dagger nematode, Xiphinema diversicaudatum, in a non-persistent manner (Valdez et al. 1974; McNamara and Eppler 1989; Hay and Close 1992; Brown and Trudgill 1998). Adults and juveniles of X. diversicaudatum transmit ArMV with equal efficiency and remain infective for up to 44 and 36 weeks, respectively, in the absence of host plants. Infectivity is retained by nematodes for periods of up to 8 months (Harrison and Winslow 1961; Eppler 1983). The virus is not retained through the molt between stages of the life cycle and is not passed from the female to the egg.

H

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Humulus lupulus (Hop)

Detailed studies on the acquisition time necessary for nematodes to become viruliferous with ArMV-H are lacking in hop. ArMV-H is also transmissible mechanically. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Cherry leaf roll virus Taxonomic position Genus: Nepovirus

(CLRV)

Family: Secoviridae

CLRV is known to cause symptomless infections in Humulus lupulus in the UK (Clark 1975; Pethybridge et al. 2008). The virus is transmitted by nematode vectors in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and by grafting. For more details of CLRV, refer to Prunus avium.

Citrus bark cracking viroid Taxonomic position Genus: Cocadviroid

(CBCVd)

Family: Pospiviroidae

CBCVd infection in plants of Humulus lupulus was reported from Slovenia (Jakse et al. 2015). The viroid-infected hop plants exhibit stunting, resulting from a shortening of the internodes of main and lateral branches, leaf yellowing and downward curling, and reduced cone production. The first symptoms appear 4–12 months after infection and plants die within 3–5 years. The viroid is mechanically sap-transmissible and also transmitted by grafting. For more details of CBCVd, refer to Citrus spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Humulus lupulus was reported from Romania, Germany, and the UK (Schmidt 1967; Schmidt and Karl 1967; Macovei 1988; Pethybridge et al. 2008). The virus-infected hop plants exhibit leaf distortion and chlorotic spots in cv. Huller Bitterer; mottled foliage and shortened internodes were also observed in some cultivars. The virus is transmitted by a number of aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Hop latent viroid Taxonomic position Genus: Cocadviroid

(HpLVd)

Family: Pospiviroidae

Humulus lupulus (Hop)

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Geographical distribution HpLVd infection was first reported in plants of commercial hops (Humulus lupulus) from Europe (Pallas et al. 1987; Puchta et al. 1988) and was found to be spread widely among the world hop-growing areas: China, Japan, Washington, Germany, New Zealand, Korea, Poland, Slovenia, the Czech Republic, Brazil, Australia, Belgium, Turkey, the USA, and the UK (Hay 1989; Barbara et al. 1990a, b; Lee et al. 1990; Adams et al. 1991, 1992; Fonseca et al. 1993; Morton et al. 1993; Matousek et al. 1994; Solarksa et al. 1995; Nelson et al. 1997; Knapic and Javornik 1998; Knabel Von et al. 1999; Barbara and Adams 2003; Pethybridge et al. 2008; Lu et al. 2012; Ziegler et al. 2014; De Jonghe et al. 2016; Gok Guler and Onelge 2017). Symptoms and host(s) HpLVd infection is latent in nature in the majority of hop cultivars (Puchta et al. 1988, 1989). Symptoms caused by infection with HpLVd have been described in cv. Omega, including chlorosis, slow growth, and the production of fewer laterals with smaller cones (Barbara et al. 1990a; Adams et al. 1991; Patzak et al. 2001). HpLVd has a very narrow host range limited primarily to Humulus lupulus, H. japonicus Sieb. and Zucc., and stinging nettle (Urtica dioica L.) in Europe (Knabel Von et al. 1999). Transmission The worldwide distribution of HpLVd is probably due to its efficient mechanical transmission and dissemination in vegetative propagules. Low transmission efficiency has been reported through pollen transfer or by seed (Matousek and Patzak 2000). Etiology and genome properties The genome consists of a single-stranded circular RNA of 253–256 nt, which is non-coding. The reference sequence is of 256 nt (X07397 = NC_003611) (Puchta et al. 1988; Hataya et al. 1992; GoraSochacka 2004; Liu et al. 2008; Giguere et al. 2014).

Hop latent virus Taxonomic position Genus: Carlavirus

(HpLV)

Family: Betaflexiviridae

Geographical distribution HpLV was first reported in plants of Humulus lupulus from Germany by Schmidt et al. (1966). HpLV is widely spread in hop gardens in Europe, the USA, Poland, China, New Zealand, South Africa, Japan, and Australia (Thresh 1969; Probasco and Skotland 1978; Adams and Barbara 1982; Adams et al. 1982; Munro 1987; Yu and Liu 1987; Von Weschmar et al. 1989; Hay et al. 1992; Kanno et al. 1993; Klein and Husfloen 1995; Pethybridge et al. 2008, 2009; Ziegler et al. 2014). Symptoms and host(s) HpLV has only been found in H. lupulus (Adams and Barbara 1982), with most cultivars remaining symptomless, although sporadic development of systemic chlorotic flecking in cv. Hersbrucker Spat has been described.

H

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Humulus lupulus (Hop)

Transmission The virus is transmitted by aphid vectors, Phorodon humuli, Myzus persicae, and Macrosiphum euphorbiae, in a non-persistent manner (Paine 1953; Adams and Barbara 1982; Eppler 1994; Crowle et al. 2006). The virus is mechanically sap-transmissible to 11 host plant species in 4 families. The virus is not seed-transmitted. Use of infected planting material (shoots) is the primary source of infection. Vegetative propagation of virus-infected plants is a major route of virus dissemination. Virion properties and genome The virions are flexuous filaments about 675 nm in length and 14 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8612 nt (AB032469 = NC_002552) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Adams and Barbara 1982; Hataya et al. 2000; Adams et al. 2004).

Hop mosaic virus Taxonomic position Genus: Carlavirus

(HpMV)

Family: Betaflexiviridae

Geographical distribution HpMV infection in plants of Humulus lupulus was first described by Salmon (1923) from the UK. HpMV is worldwide in distribution, having been described in Europe, Australia, China, South Africa, Japan, the USA, New Zealand, and North America (Mackenzie et al. 1929; Hoerner 1949; Bock 1967a; Schmelzer and Schmidt 1977; Adams et al. 1982; Yu and Liu 1987; Von Weschmar et al. 1989; Hay et al. 1992; Kanno et al. 1994; Klein and Husfloen 1995, 1995; Polak 1996; Pethybridge et al. 2000a, 2008, 2009). Symptoms and host(s) HpMV infection of most modern hop cultivars is generally symptomless. In other hop cultivars, HpMVinfected plants show initial symptoms of vein-clearing and pale green to yellow spots near veins, then leaf margins twist downward, and leaves become yellow-green, brittle, and distorted. Plants become bushy in appearance due to shortened internodes, finally leading to early death due to gradual root decline (Legg and Ormerod 1959; Thresh and Adams 1983; Probasco and Murphey 1996). Transmission The virus is transmitted through aphid vectors such as Phorodon humuli, Macrosiphum euphorbiae, and Myzus persicae in a non-persistent manner (Paine 1953; Paine and Legg 1953; Adams and Barbara 1982; Eppler 1983, 1994; Crowle et al. 2006). The virus is mechanically sap-transmissible and was not transmissible through hop seed (Legg 1965). Use of infected planting stock (shoots) acts as the primary source of spread. Virion properties and genome The virions are flexuous filaments about 650 nm in length and 14 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8550 nt (EU527979 = NC_010538) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Adams and Barbara 1980; Hataya et al. 2001; Adams et al. 2004; Poke 2008).

Humulus lupulus (Hop)

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Hop stunt viroid

(HpSVd)

Taxonomic position Genus: Hostuviroid

Family: Pospiviroidae

Geographical distribution HpSVd was first identified in Japan from plants of Humulus lupulus by Sasaki and Shikata (1977). The viroid infection on hop plants was reported from Japan, Korea, China, Europe, Asia, Africa, Oceania, North America, Germany, Slovenia, Iran, Israel, Jordan, and South America (Yamamoto et al. 1970, 1973; Sasaki and Shikata 1980; Shikata 1987; Koltunow et al. 1988; Lee and Choi 1988; Sano and Shikata 1989; Barbara et al. 1990b; Lee et al. 1990; Sano 2003, 2013; Eastwell and Nelson 2007; Guo et al. 2008; Lu et al. 2012; Radisek et al. 2012; Seigner et al. 2014). Symptoms and host(s) The viroid-infected hop plants typically have shortened internodes on the main and lateral bines, reducing plant height to approximately 3 m. Upper leaves appear curled, smaller in size, and chlorotic. Foliar symptoms of HpSVd may vary. In Japan, infected leaves droop from the base creating a characteristic X formation with laterals. Other foliar symptoms associated with HpSVd infection are a yellow-green color on the basal foliage early in the season, while yellow speckling along the major veins may also be present. Stunting appears 3–5 years after established plants become infected (Eastwell and Sano 2009; Guo et al. 2008). HpSVd has very wide host range of very economically important crops and causes various specific disorders like stunting in hop, dapple fruit on peach and plum, fruit deformation and rugosity on apricot and plum, pale fruits on cucumber, and cachexia on citrus. On the other hand, HpSVd is symptomless in grapevine and almond. Transmission The primary mode of transmission of HpSVd is mechanically through sap and also by contaminated tools (Sasaki et al. 1989; Yamamoto et al. 1973). No insect vector or seed transmission is reported. Use of shoots from the infected plants for propagation is the primary cause of this viroid spread. Etiology and genome properties The genome consists of covalently closed circular single-stranded RNA composed of 295–303 nt; the reference sequence has 302 nucleotides (X06719 = NC_001351) (Ohno et al. 1983; Momma and Takahashi 1984; Sano and Shikata 1989). The genome is highly variable, and more than 250 sequence variants are deposited in the Subviral RNA database (Lee et al. 1988; Amari et al. 2001; Gora-Sochacka 2004; Rocheleau and Pelchat 2006; Flores et al. 2011; Giguere et al. 2014).

Petunia asteroid mosaic virus Taxonomic position Genus: Tombusvirus

(PetAMV)

Family: Tombusviridae

PetAMV infection in plants of Humulus lupulus was reported from the former Czechoslovakia (Polak and Svoboda 1988; Smith et al. 1988; Pethybridge et al. 2008). The virus-infected hop plants exhibit deformed asymmetric leaves with necrotic crinkles, ringspots, line patterns, chlorotic spotting, perforated leaves, and general chlorosis. No vector is reported for this virus. The virus is transmissible by

H

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Humulus lupulus (Hop)

mechanical sap-inoculation to species from several plant families and also by grafting. For more details of PetAMV, refer to Petunia spp.

Prunus necrotic ringspot virus Taxonomic position Genus: Ilarvirus

(PNRSV)

Family: Bromoviridae

PNRSV has been reported infecting plants of Humulus lupulus worldwide, including the USA, the UK, Japan, Australia, New Zealand, the Czech Republic, Germany, Belgium, France, Romania, Yugoslavia, China, and South Africa (Wade 1962; Bock 1966, 1967b; Thresh and Ormerod 1973; Eppler and Sander 1981; Neve and Thresh 1984; Yu and Liu 1987; Dolinar 1988; Eppler 1989; Macovei 1988; Thresh et al. 1988; Von Weschmar et al. 1989; Hay et al. 1992; Johnstone et al. 1995; Klein and Husfloen 1995; Pethybridge et al. 2000b). In the USA, two isolates of PNRSV, viz., HP-1 and HP-2, are identified from hop plants (Smith and Skotland 1986). PNRSV infection on hop plants causes ring and band pattern mosaic and the necrotic crinkle mosaic symptoms (Schmelzer and Schmidt 1977). The ringspot symptoms are yellow-green, and are irregularly distributed on the leaf. Necrotic strains cause death of tissues and plant parts. In midsummer, symptoms on less severely infected plants will disappear. PNRSV in some hop cultivars also causes symptoms of necrotic crinkling of leaves which are small twisted downward and distorted. The tissue surrounding leaf veins first becomes yellow then dark brown and necrotic. The tissue area surrounding the necrosis continues to grow, and causes the characteristic crinkle symptom. Necrotic spots also appear along the veins, whereas cones remain underdeveloped and distorted. No vector transmission of PNRSV in hop is reported (Eppler and Dahdahbiglou 1991). Mechanical transmissions of PNRSV by cultural operations such as mowing, stringing, training, leaf stripping, and thinning have also been suggested (Eppler 1983; Thresh et al. 1988). Root grafting is also a possibility, considering the large root mass of commercial hop plants, its perennial nature, and close planting spaces. Glasshouse trials have indicated that PNRSV is transmissible by contact and simulated slashing between infected and virus-free plants. In some cultivars up to 51% of seeds have been infected by PNRSV (Thresh 1981). Use of infected planting material is the main source of virus spread. For more details of PNRSV, refer to Prunus persica.

Raspberry bushy dwarf virus Taxonomic position Genus: Idaeovirus

(RBDV)

Family: Unassigned

RBDV infection in plants of Humulus lupulus was reported from the UK, as a minor virus without symptom production (Pethybridge et al. 2008). The virus is mechanically sap-transmissible. For more details of RBDV, refer to Rubus spp.

Strawberry latent ringspot virus Taxonomic position Genus: Unassigned

(SLRSV)

Family: Secoviridae

Humulus lupulus (Hop)

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SLRSV infection in plants of Humulus lupulus was reported from the former Czechoslovakia (Polak and Svoboda 1988; Pethybridge et al. 2008). The virus is transmitted by the nematode vector Xiphinema diversicaudatum. The virus is mechanically sap-transmissible and also by grafting. Use of infected planting material is the primary mode of spread. For more details of SLRSV, refer to Fragaria spp.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV has been reported affecting plants of Humulus lupulus in China, as a minor latent virus (Adams 1973; Xie and Tian 1984; Pethybridge et al. 2008). There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco necrosis virus

(TNV)

Taxonomic position TNV is a tentative member of the genus Alphanecrovirus and family Tombusviridae. TNV infection in plants of Humulus lupulus was reported from Romania, France, and the former Czechoslovakia (Albrechtova et al. 1979; Chod et al. 1979; Eppler 1989; Macovei 1988; Pethybridge et al. 2008). Symptoms of infection in hop were necrotic local lesions. The virus(es) is/are mechanically sap-transmissible to a large number of hosts, and on Phaseolus vulgaris distortion, mottling and marginal chlorosis were observed, while Datura stramonium plants also become distorted, but systemic infection was unpredictable (Albrechtova et al. 1979). The virus(es) is/are transmitted by a fungal vector Olpidium brassicae in a non-persistent manner. Although the isolate of Albrechtova et al. (1979) was identified using antiserum to TNV-D (Betanecrovirus), it is not clear whether other hop isolates belong to TNV-A (Alphanecrovirus) or TNV-D. For more details of TNV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV was reported from plants of Humulus lupulus as a minor problem in Germany, causing ringspot symptoms (Adams 1972; Pethybridge et al. 2008). The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

H

1214

Humulus lupulus (Hop)

Tomato bushy stunt virus Taxonomic position Genus: Tombusvirus

(TBSV)

Family: Tombusviridae

TBSV infection in plants of Humulus lupulus was reported from Czechoslovakia (Novak and Lanzova 1976). The virus is transmissible by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants. There is no known vector for this virus. For more details of TBSV, refer to Solanum lycopersicum.

References Adams AN (1972) Virus diseases of hop production of virus-free hop plants. Rep East Malling Res Stn 1971:123–124 Adams AN (1973) Viruses latent in hop (Humulus lupulus L.) and techniques for obtaining virus-free clones. In: Proceedings of the 7th British insecticide and fungicide conference, pp 431–435 Adams AN, Barbara DJ (1980) Host range, purification, and some properties of Hop mosaic virus. Ann Appl Biol 96:201–208 Adams AN, Barbara DJ (1982) Host range, purification and some properties of two Carlaviruses from hop (Humulus lupulus): hop latent and American hop latent. Ann Appl Biol 101:483–494 Adams AN, Barbara DJ, Clark MF (1982) The occurrence and some properties of three Carlaviruses infecting hop plants (Humulus lupulus) in England. Acta Hortic 130:243–248 Adams AN, Barbara DJ, Clark MF, Davies DL (1986) Association of arabis mosaic virus with hop chlorotic disease. Acta Hortic 193:59–60 Adams AN, Bardara MF, Clark DF, Davies DL (1987) The etiology of hop chlorotic disease. Ann Appl Biol 111:365–371 Adams AN, Barbara DJ, Morton A (1991) Effects of Hop latent viroid on weight and quality of the cones of the hop cultivar Wye challenger. Ann Appl Biol 118:126–127 Adams AN, Morton A, Barbara DJ, Ridout MS (1992) The distribution and spread of Hop latent viroid within two commercial plantings of hop (Humulus lupulus). Ann Appl Biol 121:585–592 Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Albrechtova L, Chod J, Kriz J, Fencl J (1979) Detection of Apple mosaic virus and Tobacco necrosis virus in Hop. Phytopathol Z 94:45–55 Amari K, Gomez G, Myrta A, Bdi T, Pallás V (2001) The molecular characterization of 16 new sequence variants of Hop stunt viroid reveals the existence of invariable regions and a conserved hammerhead-like structure on the viroid molecule. J Gen Virol 82(4):953–962 Barbara DJ, Adams AN (2003) Hop latent viroid. In: Hadidi A, Flores R, Randles JW, Semancik JS (eds) Viroids. CSIRO Press, Collingwood, pp 213–217 Barbara DJ, Morton A, Adams AN, Green CP (1990a) Some effects of Hop latent viroid on two cultivars of hop (Humulus lupulus) in the UK. Ann Appl Biol 117:359–366 Barbara DJ, Morton A, Adams AN (1990b) Assessment of UK hops for the occurrence of hop latent and hop stunt viroids. Ann Appl Biol 116:265–272 Bock KR (1966) Arabis mosaic and Prunus necrotic ringspot viruses in hop (Humulus lupulus L.). Ann Appl Biol 57:131–140 Bock KR (1967a) Hop mosaic virus. Rep East Malling Res Stn 1966:163–165 Bock KR (1967b) Strains of Prunus necrotic ringspot virus in hop (Humulus lupulus L.). Ann Appl Biol 59:437–446 Brown DJF, Trudgill DL (1998) Nematode transmission of plant viruses: a 30 year perspective. Host pathogen interactions and crop protection. SCRI Annual Report, pp 121–125 Cartledge EG (1956) An investigation into the diseases of Tasmanian hops. Tas J Agric 27:210–219 Chod J, Jokes M, Novak M (1979) The electron-microscopic proof of Tobacco necrosis virus in hop plants. Biol Plant 21:152–153 Clark MF (1975) Virus diseases of hop. Rep East Malling Res Stn 1974:124 Crowle DR, Pethybridge SJ, Leggett GW, Sherriff LJ, Wilson CR (2003) Diversity of the coat protein-coding region among Ilarvirus isolates infecting hop in Australia. Plant Pathol 52:655–662

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Crowle DR, Pethybridge SJ, Wilson CR (2006) Transmission of hop latent and hop mosaic carlaviruses by Macrosiphum euphorbiae and Myzus persicae. J Phytopathol 154:745–747 Davies DL, Clark MF (1983) A satellite-like nucleic acid of Arabis mosaic virus associated with hop nettlehead disease. Ann Appl Biol 103:439–448 De Jonghe K, Van Bogaert N, Vandierendonck S, Smagghe G, Maes M (2016) First report of Hop latent viroid in Belgian hops. Plant Dis 100:1956 Dolinar M (1988) Ilar-viruses in hop gardens in Slovenia, Yugoslavia. In: Eppler A (ed) Proceedings of the international workshop on hop virus diseases, Rauischholzhausen Castle, pp 83–88 Eastwell KC, Druffel KL (2012) Complete genome organization of American hop latent virus and its relationship to carlaviruses. Arch Virol 157:1403–1406 Eastwell KC, Nelson ME (2007) Occurrence of viroids in commercial hop (Humulus lupulus L.) production areas of Washington State. Online. Plant Health Prog. https://doi.org/10.1094/PHP-2007-1127-01-RS Eastwell K, Sano T (2009) Hop stunt disease (HSVd). In: The compendium of hop diseases, arthropod pests and other disorders. APS Press, St. Paul, USA. pp 48–51. 93 page. ISBN 978-0-89054-376-4 Eppler A (1983) Transmission of hop viruses and the role of wild and escaped hops as sources of virus spread. Medelingen van de Faculteit Landbouwwetenschappen Rijk’lunivesitent Gent 48:883–892 Eppler A (1989) Presence of viruses in the hop growing region of Alsace (France). In: Eppler A (ed) Proceedings of the international workshop on hop virus diseases, Rauischholzhausen Castle, p 209 Eppler A (1994) Ecology of aphids on hops and its significance on spread of hop viruses. Z Pflanzenkr Pflanzenschutz 102:2–15 Eppler A, Dahdahbiglou R (1991) Experiments on the non-vectorial transmission of PNRV in hops. Med Fac Landbouww Rijksuniv Gent 56:577–587 Eppler A, Sander E (1981) The infection of German hops with Prunus necrotic ringspot virus. Medelingen van de Faculteit Landbouwwetenschappen Rijksunivesitent Gent 46:1115–1121 Flores R, Daros J, Hernandez J-A, Di Serio F (2011) Viroids. In: Encyclopedia of life sciences (ELS). John Wiley and Sons, Ltd: Chichester. https://doi.org/10.1002/9780470015902.a0000434.pub3 Fonseca MEN, Marinho VLA, Nagata T (1993) Hop latent viroid in hop germ plasm introduced into Brazil from the United States. Plant Dis 77:952 Giguere T, Raj Adkar-Purushothama C, Perreault J-P (2014) Comprehensive secondary structure elucidation of four genera of the family Pospiviroidae. PLoS One 9(6):e98655. https://doi.org/10.1371/journal.pone.0098655 Gok Guler P, Onelge N (2017) First report of Hop latent viroid in Turkey. J Plant Pathol 99:799–818 Gora-Sochacka A (2004) Viroids: unusual small pathogenic RNAs. Acta Biochim Pol 51(3):587–607 Guo L, Liu S, Wu Z, Mu L, Xiang B, Li S (2008) Hop stunt viroid (HSVd) newly reported from hop in Xinjiang, China. Plant Pathol 57:764 Harrison BD, Winslow RD (1961) Laboratory and field studies on the relation of ArMV to its nematode vector Xiphinema diversicaudatum. Ann Appl Biol 49:621–633 Hataya T, Hikage K, Sunda N, Nagata T, Li S, Itoga Y, Tanikoshi T, Shikata E (1992) Detection of Hop latent viroid (HLVd) using reverse transcription and polymerase chain reaction (RT-PCR). Ann Phytopathol Soc Jpn 58:677–684 Hataya T, Uchino K, Arimoto R, Suda N, Sano T, Shikata E, Uyeda I (2000) Molecular characterisation of Hop latent virus and phylogenic relationships among viruses closely related to carlaviruses. Arch Virol 145:2503–2524 Hataya T, Arimoto R, Suda N, Uyeda I (2001) Molecular characterisation of Hop mosaic virus: its serological and molecular relationships to Hop latent virus. Arch Virol 146:1935–1948 Hay FS (1989) Studies on the viruses of hop (Humulus lupulus L.) in New Zealand. PhD thesis, Lincoln University, Christchurch, New Zealand Hay FS, Close RC (1992) Distribution of Xiphinema diversicaudatum (Micoletzky, 1927) Thorne, 1939 in commercial hop (Humulus lupulus L.) gardens in New Zealand and implications for the spread of arabis mosaic virus. NZJ Crop Hortic Sci 20:367–370 Hay FS, Close RC, Fletcher JD, Ashby JW (1992) Incidence and spread of viruses in hop (Humulus lupulus L.) in New Zealand. NZJ Crop Hortic Sci 20:319–327 Hoerner GR (1949) Hop diseases in the United States. Brewers’ Dig 24:45–51 Ito T, Yoshida K (1998) Reproduction of apple fruit crinkle disease symptoms by apple fruit crinkle viroid. Acta Hortic 472:587–594 Ito T, Kanematsu S, Koganezawa H, Tsuchizaki T, Yoshida Y (1993) Detection of a viroid associated with apple fruit crinkle disease. Ann Phytopathol Soc Jpn 59:520–527 Jakse J, Radisek S, Pokorn T, Matousek J, Javornik B (2015) Deep-sequencing revealed citrus bark cracking viroid (CBCVd) as a highly aggressive pathogen on hop. Plant Pathol 64:831–842 Johnstone GR, Munro D, Brown GS, Skotland CB (1995) Serological detection, occurrence and spread of ilarviruses in temperate fruit crops, hops and roses in Tasmania. Acta Hortic 386:132–135 Kanno Y, Yoshikawa N, Takahashi T (1993) Some properties of hop latent and apple mosaic viruses isolated from hop plants and their distributions in Japan. Ann Phytopathol Soc Jpn 59:651–658

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Kanno Y, Iida H, Yoshikawa N, Takahashi T (1994) Some properties of Hop mosaic virus isolated in Japan. Ann Phytopathol Soc Jpn 60:675–680 Keyworth WG, Davies DLG (1946) Nettlehead disease of the hop (Humulus lupulus). J Pomol Hortic Sci 22:134–139 Klein RE, Husfloen SD (1995) Incidence of five viruses in hop (Humulus lupulus) in the Pacific Northwest. Plant Dis 79:425–428 Knabel Von S, Seigner L, Wallnofer PR (1999) Detection of Hop latent viroid (HLVd) using the polymerase chain reaction (PCR). Gesunde Pflanzen 51:234–239 Knapic V, Javornik B (1998) Viroidi-Povrocitelji Rastlinskih Bolezni. Sodobno Kmetijstvo 31:462–465 Koltunow AM, Krake LR, Rezaian MA (1988) Hop stunt viroid in Australia grapevine cultivars: potential for hop infection. Aust Plant Pathol 17:7–9 Kominek P, Svoboda P, Abou Ghanem-Sabanadzovic N (2003) Improved detection of arabis mosaic virus in grapevine and hop plants. Acta Virol 47:199–200 Lee JY, Choi JK (1988) Detection of hop stunt viroid in Korea. Korean J Plant Pathol 4(3):234–237 Lee JY, Puchta H, Ramm K, Sanger HL (1988) Nucleotide sequence of the Korean strain of Hop stunt viroid (HSV). Nucleic Acids Res 16:8708 Lee JY, Lee SH, Sanger HL (1990) Viroid diseases occurring on Korean hop plants. Korean J Plant Pathol 6:256–260 Legg JT (1965) Mechanical transmission of Hop mosaic virus. Nature 208:1017–1018 Legg JT, Ormerod PJ (1959) Research on hop virus diseases at East Malling. Rep East Malling Res Stn 1958:157–160 Liu S, Li S, Zhu J, Xiang B, Cao L (2008) First report of Hop latent viroid (HLVd) in China. Plant Pathol 57:400 Lu W, Zhang Z, Xu P, Liu S, Wang H, Jiang D, Li S (2012) Simultaneous detection of three viroid species infecting hops in China by multiplex RT-PCR. J Phytopathol 160:308–310 Mackenzie D, Salmon ES, Ware WM, Williams R (1929) The mosaic disease of the hop. Ann Appl Biol 16:359–381 Macovei A (1988) A survey of the Romanian virus diseases and the electronmicroscopial characterization of the viruses involved. In: Proceedings of the international workshop on hop virus diseases, Giessen, pp 19–22 Matousek J, Patzak J (2000) A low transmissibility of Hop latent viroid (HLVd) through a generative phase of hop (Humulus lupulus L.). Biol Plant 43:145–148 Matousek J, Trnena L, Svoboda P, Ruzkova P (1994) Analysis of Hop latent viroid (HLVd) in commercial hop clones in Czech Republic. Rost Vyr 40:973–983 McNamara DG, Eppler A (1989) Arabis mosaic virus and Xiphinema diversicaudatum in German hops. In: Proceedings of the international workshop on hop virus diseases, Giessen, p 198 Momma T, Takahashi T (1984) Developmental morphology of Hop stunt viroid-infected hop plants and analysis of their cone yield. J Phytopathol 110:1–4 Morton A, Barbara DJ, Adams AN (1993) The distribution of Hop latent viroid within plants of Humulus lupulus and attempts to obtain viroid-free plants. Ann Appl Biol 123:47–53 Munro D (1987) Viruses infecting hop, Humulus lupulus L. in Australia. Aust J Agric Res 38:83–90 Nelson ME, Klein RE, Skrzeczkowski LJ (1997) Occurrence of Hop latent viroid (HLVd) in hops in Washington State. (Abstr.). Phytopathology 88:S108 Neve RA, Thresh JM (1984) Prunus necrotic ringspot virus in hops. Eng Hops 1:6–7 Novak JB, Lanzova J (1976) Identification of Alfalfa mosaic virus and Tomato bushy stunt virus in hop (Humulus lupulus L.) and grapevine (Vitis vinifera subsp. sativa (DC/Hegi) plants in Czeschoslovakia). Biol Plant 18:152–154 Ohno T, Takamatsu M, Meshi T, Okada Y (1983) Hop stunt viroid: molecular cloning and nucleotide sequence of the complete cDNA copy. Nucleic Acids Res 11:6185–6197 Paine J (1953) Insect vector studies with mosaic and other virus diseases of the hop. Rep East Malling Res Stn 1952:120–123 Paine J, Legg JT (1953) Transmission of hop mosaic by Phorodon humuli (Schrank). Nature 171:263–264 Pallas V, Navarro A, Flores R (1987) Isolation of a viroid-like RNA from hop different from Hop stunt viroid. J Gen Virol 68:3201–3205 Patzak J, Matousek J, Krofta K, Svoboda P (2001) Hop latent viroid (HLVd)-caused pathogenesis: effects of HLVd infection on lupulin composition of meristem culture-derived Humulus lupulus. Biol Plant 44:579–585 Pethybridge SJ, Wilson CR, Sherriff LJ, Leggett GW, Munro D (2000a) Virus incidence in Australian hop (Humulus lupulus L.) gardens and cultivar differences in susceptibility to infection. Aust J Agric Res 51:685–689 Pethybridge SJ, Wilson CR, Leggett GW (2000b) Spread of Prunus necrotic ringspot virus in Australian hop gardens. Phytopathology 90(6 Suppl):S60–S61 E Pethybridge SJ, Wilson CR, Hay FS, Leggett GW, Sherriff LJ (2002a) Mechanical transmission of Apple mosaic virus in Australian hop (Humulus lupulus) gardens. Ann Appl Biol 141:77–85 Pethybridge SJ, Wilson CR, Hay FS, Leggett GW, Sherriff LJ (2002b) Effect on viruses on agronomic and brewing characteristics of four hop (Humulus lupulus) cultivars in Australia. Ann Appl Biol 140:97–105

Humulus lupulus (Hop)

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Pethybridge SJ, Hay FS, Barbara DJ, Eastwell KC, Wilson CR (2008) Viruses and viroids infecting hop: significance, epidemiology, and management. Plant Dis 92:324–338 Pethybridge SJ, Fletcher JD, Hay FS, Beatson RA (2009) Prevalence and incidence of viruses in New Zealand hop (Humulus Iupulus) gardens. NZJ Crop Hortic Sci 37(3):235–241 Poke FS (2008) Hop mosaic virus: complete nucleotide sequence and relationship to other carlaviruses. Arch Virol 153(8):1615–1619 Polak J (1996) Distribution of Hop mosaic virus in hop gardens of the Czech Republic. Ochr Rostl 32:9–14 Polak J, Svoboda P (1988) The first results of screening hop clones and varieties for the presence of some viruses in Czechoslovakia using ELISA. In: Eppler A (ed) Proceedings of the international workshop on hop virus diseases, Rauischholzhausen Castle, pp 27–32 Probasco EG, Murphey JM (1996) The effects of hop viruses on brewing and agronomic characteristics in the hop variety Chinook. MBAA Tech Q 33:160–165 Probasco EG, Skotland CB (1976a) A technique for the differential isolation of hop mosaic and Hop latent virus. Can J Microbiol 22:1160–1162 Probasco EG, Skotland CB (1976b) Host range and properties of the third rodshaped virus found in hops. Proc Am Phytopathol Soc 3:319 Probasco EG, Skotland CB (1978) Host range, general properties, purification and electron microscopy of Hop latent virus. Phytopathology 68:277–281 Puchta H, Ramm K, Sanger HL (1988) The molecular structure of Hop latent viroid (HLV), a new viroid occurring worldwide in hops. Nucleic Acids Res 16:4197–4216 Puchta H, Ramm K, Sanger HL (1989) Hop latent viroid (HLVd), a new viroid occurring worldwide in hops. In: Proceedings of the international workshop on hop virus diseases, Giessen, pp 181–191 Radisek S, Majer A, Jakse A, Javornik B, Matoušek J (2012) First report of hop stunt viroid infecting hop in Slovenia. Plant Dis 96:592 Rocheleau L, Pelchat M (2006) The subviral RNA database: a toolbox for viroids, the hepatitis delta virus and satellite RNAs research. BMC Microbiol 6:24 Salmon ES (1923) The mosaic disease of the hop. J Minist Agric 29:1–7 Sano T (2003) Hop stunt viroid. In: Hadidi A, Flores R, Randles JW, Semancik JS (eds) Viroids. CSIRO Publishing, Collingwood, pp 207–212 Sano T (2013) History, origin, and discovery of hop stunt disease and hop stunt viroid. Acta Hortic 1010:87–96 Sano T, Shikata E (1989) Hop stunt disease. In: Proceedings of the international workshop on hop virus diseases, Giessen, pp 159–164 Sano T, Sasaki M, Shikata E (1985) Apple mosaic virus isolated from hop plants in Japan. Ann Appl Biol 106:305–312 Sano T, Yoshida H, Goshono M, Monma T, Kawasaki H, Ishizaki K (2004) Characterization of a new viroid strain from hops: evidence for viroid speciation by isolation in different host species. J Gen Plant Pathol 70:181–187 Sano T, Isono S, Matsuki K, Kawaguchi-Ito Y, Tanaka K, Kondo K, Iijima A, Bar-Joseph M (2008) Vegetative propagation and its possible role as a genetic bottleneck in the shaping of the Apple fruit crinkle viroid populations in apple and hop plants. Virus Genes 37:298–303 Sasaki M, Shikata E (1977) On some properties of hop stunt disease agent, a viroid. Proc Jpn Acad 55B:109–112 Sasaki M, Shikata E (1980) Hop stunt disease, a new viroid disease occurring in Japan. Rev Plant Protect Res 13:97–109 Sasaki M, Fukamizu K, Yamamoto K, Ozawa T, Kurokawa M, Kagami Y (1989) Epidemiology and control of hop stunt disease. In: Eppler A (ed) Proceedings of the international workshop on hop virus diseases, Rauischholzhausen 1988. Deutsche Phytomedizinische Gesellschaft, Ulmer Verlag, Stuttgart, pp 165–178 Schmelzer K, Schmidt HE (1977) Spezialkulturen. In: Klinkowski M et al (eds) Pflanzliche Virologie, vol 3, Third edn. Academie, Verlag, pp 294–364 Schmidt HE (1967) New results of hop virus research. In: Proceedings of the 6th conference of the czechoslovak plant virologists, pp 47–53 Schmidt HE, Karl E (1967) Untersuchungen uber eine Scheckung am Japanischem Hopfen Humulus scandens (Lour.) Merr. Phytopathol Z 56:272–278 Schmidt HE, Schmidt HB, Eisbein K (1966) Die mechanische ubertragung eines stabschenformingen hopfenvirus auf krautige testpflanzen. Zentbl Bakt Parasitkore 120:461–466 Seigner L, Lutz A, Seigner E (2014) Monitoring of dangerous hop viroids and viruses in German hop production. JuliusKuhn-Archiv 447:115–116 Shikata E (1987) Hop stunt. The viroids. TO Diener, Plenum Press, New York, 279–290. Smith DR, Skotland CB (1986) Host range and serology of Prunus necrotic ringspot virus serotypes isolated from hops (Humulus lupulus) in Washington. Plant Dis 70:1019–1023

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Smith KM, Dunez J, Lelliott RA, Phillips DH, Archer SA (eds) (1988) European handbook of plant diseases. Blackwell Scientific Publications, Oxford Solarksa E, Skomra U, Kitlinksa J, Wojcierowski J (1995) The occurrence of Hop latent viroid (HLVd) in hop plants in Poland. Phytopathol Polonica 10:55–59 Thresh JM (1969) Hop latent virus. Rep East Malling Res Stn 1968:41 Thresh JM (1981) Virus diseases of hop. Rep East Malling Res Stn 1980:85–86 Thresh JM, Adams AN (1983) Hop mosaic disease. Rep East Malling Res Stn 1982:173–175 Thresh JM, Edwards P (1983) Virus and virus diseases of hop – Arabis mosaic virus (AMV). Rep East Malling Res Stn 1982:89 Thresh JM, Ormerod PJ (1973) Virus diseases of hop – spread of Prunus necrotic ringspot virus in hop. Rep. East Malling Res Stn 1972:163–164 Thresh JM, Ormerod PJ (1989) Arabis mosaic virus in English hop plantings. In: Proceedings of the international workshop on hop virus diseases, Giessen, pp 43–54 Thresh JM, Pitcher RS (1978) The spread of nettlehead and related virus diseases of hop. In: Scott PR, Bainbridge A (eds) Plant disease epidemiology. Blackwell, Oxford, pp 291–298 Thresh JM, Pitcher RS, McNamara DG, Ormerod PJ (1972) The spread and control of nettlehead and related diseases of hop. Rep East Malling Res Stn 1971:155–162 Thresh JM, Barbara DJ, Ormerod PJ (1988) The incidence and spread of Prunus necrotic ring spot virus in English hop plantings. In: Eppler A (ed) Proceedings of the international workshop on hop virus diseases, Rauischholzhausen Castle, pp 71–82 Valdez RB, McNamara DG, Ormerod PJ, Pitcher RS, Thresh JM (1974) Transmission of the hop strain of Arabis mosaic virus by Xiphinema diversicaudatum. Ann Appl Biol 76:113–122 Von Weschmar WB, Brits G, Coleman T (1989) Viruses in hops and aspects of virus epidemiology and hop production in South Africa at 34 southern latitude. In: Proceedings of the international workshop on hop virus diseases, Giessen, pp 33–42 Wade GC (1962) Hop diseases in Tasmania. Tasmania J Agric 7:261–268 Xie H, Tian B (1984) A strain of Tobacco mosaic virus infecting hops in the Xinjiang province. In: Symposium on viruses and agriculture, China, p 7 Yamamoto H, Kagami Y, Kurokawa M, Nishimura S, Kubo S, Inoue M, Murayama D (1970) Studies on hop stunt disease I. Memories Res Fac Agric Hokkaido Univ 7:491–512 Yamamoto H, Kagami Y, Kurokawa M, Nishimura S, Ukawa S, Kubo S (1973) Studies on hop stunt disease in Japan. Rep Res Lab Kirin Brewery Co Ltd 16:49–62 Yu J, Liu Y (1987) The occurrence of three viruses in hop (Humulus lupulus L.) in China. Plant Pathol 36:38–44 Ziegler A, Kawka M, Przybys M, Doroszewska T, Skomra U, Kastirr U, Matousek J, Schubert J (2014) Detection and molecular analysis of Hop latent virus and Hop latent viroid in hop samples from Poland. J Kult 66:248–254

Hyacinthus spp. Family: Asparagaceae

Ornamental

Arabis mosaic virus

(ArMV)

Taxonomic position Genus: Nepovirus

Family: Secoviridae

ArMV infection in plants of Hyacinthus orientalis was reported from Lithuania (Europe) (Navalinskiene and Samuitiene 2006; Samuitiene et al. 2008). The leaves of the virus-infected hyacinth plants show chlorotic or pale brown necrotic stripes and spots of various shapes. Leaves are distorted. The virus is transmitted by a nematode vector, Xiphinema diversicaudatum, in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Hyacinthus spp.

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Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Hyacinthus spp. was reported from the UK and the Netherlands causing bright yellow foliar streaking symptoms (Ainsworth 1938; Loebenstein et al. 1995; Chen 2003). The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Hyacinth mosaic virus Taxonomic position Genus: Potyvirus

(HyaMV)

Family: Potyviridae

Geographical distribution HyMV infection in plants of Hyacinthus orientalis was first reported from Bulgaria and Oregon, USA, by Atanasoff (1928). The virus spreads in Australia, Bulgaria, the Netherlands, Lithuania, the UK, and the USA (Navalinskiene and Samuitiene 2001, 2006; Pham et al. 2011), and presumably wherever the crop is grown. Symptoms and host(s) The virus-infected hyacinth plants show a mottle on the leaves mainly on the basal parts that range in color from light green to bright yellow. The mottle is stripe-like or consists of many spots and ring patterns. Flower stalks of some cultivars show stripes and ring patterns. Flower breaking is also noticed (Derks and Vink-Van den Abeele 1980). Transmission The virus is transmitted by aphid vectors Myzus persicae and Macrosiphum euphorbiae in a non-persistent manner. The virus is also mechanically sap-transmissible, and several (three to nine) families are susceptible. Virion properties and genome The virions are non-enveloped, flexuous filaments, measuring about 740–750 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9500 nt (KY828925 = NC_037051) (Revers and Garcia 2015; Wylie et al. 2017).

Ornithogalum mosaic virus Taxonomic position Genus: Potyvirus

(OrMV)

Family: Potyviridae

A virus indistinguishable from OrMV was identified in naturally infected plants of Hyacinthus orientalis in Oregon, USA, and transmitted to Ornithogalum by the aphids Myzus persicae and

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Hyacinthus spp.

Macrosiphum euphorbiae in a non-persistent manner (Smith and Brierley 1944). The virus is also mechanically sap-transmissible. For more details of OrMV, refer to Ornithogalum spp.

Tobacco necrosis virus

(TNV)

Taxonomic position TNV is a tentative member of the genus Alphanecrovirus and family Tombusviridae TNV infection in plants of Hyacinthus orientalis was reported from Lithuania (Europe) (Asjes 1974; Derks et al. 2003; Navalinskiene and Samuitiene 2006). The leaves of the virus-infected hyacinth plants develop pale brown stripes and spots which later become necrotic. TNV infection causes distortion, narrowing of leaves. Severely infected plants do not flower. The virus is transmitted through the zoospores of the fungus Olpidium brassicae and is also mechanically sap-transmissible. It is not clear whether the TNV isolates from hyacinth are of TNV-A (Alphanecrovirus) or TNV-D (Betanecrovirus). For more details of TNV, refer to Nicotiana tabacum.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in plants of Hyacinthus orientalis was reported from Europe, in Lithuania and the Netherlands (Asjes 1974, 1994; Navalinskiene and Samuitiene 2001, 2006). The virus-infected hyacinth leaves and stalks show pale green to yellow spots and stripes. These symptoms may turn to brown or grey necrotic stripes later in the season. In some cases, leaves tear and bend along stripes. Transparent and in the later stage yellow or brown spots become visible on the bulb scales. Necrosis typical of TRV infection is seen in cross sections of infected bulbs. The virus is transmitted by Trichodorus and Paratrichodorus nematode vectors and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Hyacinthus spp. was reported from the Netherlands (Asjes 1974). The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato aspermy virus Taxonomic position Genus: Cucumovirus

(TAV)

Family: Bromoviridae

Hydrangea spp.

1221

TAV infection in plants of Hyacinthus spp. was reported from China (Shu et al. 1990). The virusinfected plants show black necrotic spots on the bulb. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of TAV, refer to Solanum lycopersicum.

References Ainsworth GC (1938) A note on certain viruses of the cucumber virus 1 type isolated from monocotyledonous plants. Ann Appl Biol 24:867–869 Asjes CJ (1974) Soil-borne viral diseases in ornamental bulbous crops and their control in the Netherlands. Agric Environ 1:303–315 Asjes CJ (1994) Occurrence of Tobacco rattle virus in ornamental bulbous crops in the Netherlands. Acta Hortic 377:349–356 Atanasoff D (1928) Mosaic disease of flower bulb plants. Bull Soc Bot Bulgarie 2:52 Chen YK (2003). Occurrence of cucumber mosaic virus in ornamental plants and perspectives of transgenic control. PhD thesis, Wageningen University, The Netherlands, p 144 Derks AFLM, Vink-Van den Abeele JL (1980) Hyacinth mosaic virus: symptoms in hyacinths, serological detection and relationships with other potyviruses. Acta Hortic 109:495–502 Derks AFLM, Pham KTK, Lemmers MEC, Blom-Barnhoorn GJ (2003) Augustaziek ook in hyacint en lelie. Bloembollen Visie 5:18–19 Loebenstein G, Lawson RH, Brunt AA (1995) Virus and virus-like diseases of bulb and flowering crops. Wiley, Chichester, p 543 Navalinskiene M, Samuitiene M (2001) Viruses affecting some bulb and corn flower crops. Biologia 4:40–42 Navalinskiene M, Samuitiene M (2006) Identification of agents associated with viral and phytoplasmal diseases in common hyacinth (Hyacinthus orientalis L). Biologija 2:54–58 Pham KTK, de Kock MJD, Lemmers MEC, Derks AFLM (2011) Molecular identification of potyviruses infecting bulbous ornamentals by the analysis of coat protein (CP) sequences. Acta Hortic 901:167–172 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Samuitiene M, Navalinskiene M, Jackeviciene E (2008) Arabis mosaic virus on ornamental plants. Biologia 54:264–268 Shu XZ, Zhu SF, Hu WZ, Liu JJ, Chen CL, Wang SQ, Chen YF (1990) Identification of tomato aspermy virus infecting chrysanthemum and hyacinth. Virol Sin 5(2):186–192 Smith FF, Brierley P (1944) Ornithogalum mosaic virus. Phytopathology 34:497–503 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Hydrangea spp. Family: Hydrangeaceae

Ornamental

Alfalfa mosaic virus

(AMV)

Taxonomic position Genus: Alfamovirus

Family: Bromoviridae

AMV was reported in plants of Hydrangea spp. from the USA and Italy (Belli 1968; Lockhart et al. 2013). The virus-infected hydrangea plants exhibit mottling, vein yellowing, yellow leaf blotching (calico symptoms), and mosaic on apical foliage (Chiko and Godkin 1986). The virus is transmitted by a number of aphid species in a non-persistent manner. It is also transmissible by mechanical

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Hydrangea spp.

sap-inoculation but not by contact between plants. The use of infected cuttings will help in the spread of the virus. For more details of AMV, refer to Medicago sativa.

Cherry leaf roll virus

(CLRV)

Taxonomic position Genus: Nepovirus

Family: Secoviridae

CLRV infection in plants of Hydrangea macrophylla was reported from New Zealand (Veerakone et al. 2012). The virus-infected hydrangea plants exhibited severe leaf deformation and chlorotic symptoms. The virus is transmitted by nematode vectors in a non-persistent manner. The virus is also transmissible by mechanical sap-inoculation, and by grafting. For more details of CLRV, refer to Prunus avium.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Hydrangea macrophylla was reported from Japan, Belgium, and Italy (Tamura and Komuro 1967; Bang et al. 2001). The virus-infected hydrangea plants exhibit stunting, leaf mosaic, and flower breaking symptoms (Bertaccini et al. 2015). The virus is transmitted by a number of aphid species in a non-persistent manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Elm mottle virus

(EMoV)

Synonyms Hydrangea mosaic virus (HdMV) Taxonomic position Genus: Ilarvirus

Family: Bromoviridae

HdMV was first reported from From West Sussex, UK, in plants of Hydrangea macrophylla (Thomas et al. 1983). The virus spreads in Italy and Southern Australia (Bertaccini et al. 2015). The virus-infected hydrangea plants exhibit distinct leaf symptoms of a mottled mosaic of light- and dark-green segments, and the flowers are not affected. The virus is mechanically sap-transmissible. No vector is known for this virus. However thrips vectors help in virus transmission by damaging the leaves while feeding and allow the virus entry into plants through a virus-carrying pollen. For more details of EMoV, refer to Ulmus spp.

Hydrangea chlorotic mottle virus Taxonomic position Genus: Carlavirus

(HCMoV)

Family: Betaflexiviridae

Hydrangea spp.

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Geographical distribution HCMoV infection in plants of Hydrangea macrophylla was reported from the USA, Korea, and New Zealand (Caballero et al. 2009; Tang et al. 2010; Song et al. 2016). Symptoms and host(s) The virus-infected H. macrophylla (var. ‘Endless Summer’) plants produce leaf blistering, reddening, chlorotic ringspots, and chlorotic netting; the virus also readily infect other macrophylla cultivars. Symptoms of chlorotic leaf mottling and distortion are also noticed in some cultivars (Tang et al. 2010). Symptoms in the cultivar ‘Endless Summer’ were more frequent during the spring but less so during the summer. HCMoV is only known to infect H. macrophylla. Transmission The virus is transmitted by the green peach aphid (Myzus persicae) in a non-persistent manner. Virion properties and genome The virions are flexuous filaments about 650 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8433 nt (NC_012869; EU754720) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Adams et al. 2004).

Hydrangea ringspot virus Taxonomic position Genus: Potexvirus

(HdRSV)

Family: Alphaflexiviridae

Geographical distribution HdRSV was first reported in plants of Hydrangea macrophylla from the USA by Brierley (1954). The virus spreads in Europe, the USA, New Zealand, Canada, Italy, Korea, and the Czech Republic (Welvaert and Samyn 1973; Allen et al. 1985; Chiko and Godkin 1986; Mertelik and Kloudova 2009; Bertaccini et al. 2015; Song et al. 2016) and also occurs wherever hydrangea are cultivated. Symptoms and host(s) The virus-infected hydrangea plants show leaves with chlorotic to brown leaf spots or rings. Leaves are sometimes crinkled or distorted, a decreased number of florets may be seen in the inflorescence is decreased, and the plants may be severely stunted. In case of H. macrophylla, the virus caused mild to severe diffuse chlorosis and diffuse chlorotic spots. Infected Hydrangea serrata plants showed unusual symptoms of conspicuous chlorotic rings, yellow blotches, and line patterns in summer (from June to August), turning red-purple later in the season (from September to November) (Mertelik and Kloudova 2011). Symptoms are often more prominent on plants in the early spring and late summer. Transmission The virus is mechanically sap-transmissible to 20 different species in 12 families. Chenopodium amaranticolor, Chenopodium quinoa, and Gomphrena globosa are local lesion hosts producing chlorotic or necrotic lesions. Virus transmission of about 12% also takes place through cutting knives while collecting the cuttings for planting (Brierley and Lorentz 1957). The virus is not known to be seed-transmitted, and no vector has been identified.

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Hydrangea spp.

Virion properties and genome The virions are flexuous filaments measuring about 470–580 nm in length and 13 nm in diameter. The genome consists of a single-stranded RNA of 6185 nt (AY707100 = NC_006943) and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type and 18–27 kDa in size (Adams et al. 2004; Hughes et al. 2005).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Hydrangea spp. was reported from the USA, Bulgaria, and Portugal (Ruter and Gitaitis 1993; Louro 1996). The virus-infected hydrangeas were symptomless or exhibited chlorotic and necrotic spot symptoms. The virus is transmitted by thrips vectors in a persistentpropagative manner, and is also transmissible by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Hydrangea spp. was reported from British Columbia (Canada) and the USA (Anderson 1958; Chiko and Godkin 1986). The virus-infected hydrangea leaves show faint chlorotic ringspots and also faint mottling. Leaf chlorosis with green blotches on leaves and stunting of plants is sometimes observed. In cultivars of Hydrangea macrophylla, symptoms of uneven chlorotic crinkling of leaves with stunted growth of plants may be observed. The virus is transmitted by nematode vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Hydrangea spp. was reported from the USA (Brierley 1954; Lawson 1995). The virus-infected hydrangea plants are stunted, and leaves are smaller than normal, often appearing narrow and irregular in shape. The leaves show a dull chlorosis with diffuse-bordered dark green blotches superimposed. Under favorable environmental conditions, the inoculated hydrangea plants express chlorotic ring and yellow vein pattern symptoms. The virus is transmitted by a nematode vector, Xiphinema americanum, in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. Vegetative propagation is the principal method of ToRSV spread in hydrangea. For more details of ToRSV, refer to Solanum lycopersicum.

Hydrangea spp.

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Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Hydrangea spp. was reported from Portugal, Bulgaria, and the USA (Allen et al. 1983, 1985; Hausbeck et al. 1992; Louro 1996; Bakardjieva et al. 1998). The virus-infected hydrangea plants express ringspot symptoms in the foliage (Allen et al. 1985). Use of vegetative planting material from infected plants is the primary source for spread of the virus. The virus is transmitted by thrips vectors in a persistent-propagative manner; mechanical sap-inoculation to Gomphrena globosa produced local lesions. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Allen TC, McMorran JP, Locatelli EA (1983) Isolation of Tomato spotted wilt virus from Hydrangea and four weed species. Plant Dis 67:429–431 Allen TC, McMorran JP, Lawson RH (1985) Detection and identification of viruses in hydrangea. Acta Hortic 164:85–90 Anderson CW (1958) Tobacco ring spot virus on florist’s Hydrangea. Plant Dis Reptr 42:932–933 Bakardjieva N, Denkova S, Hristova D (1998) Tomato spotted wilt virus on ormamental species in Bulgaria. Biotech Biotechnol Equip 12:49–52 Bang JH, Park SJ, Lee KH, Choi JK, Lee SY (2001) Characterization of cucumber mosaic virus isolated from Hydrangea macrophylla for. otaksa (Sieb. Et Zucc.) Wils. Res Plant Dis 7:1–7 Belli G (1968) Researches on a vein clearing of Hydrangea. Phytopathol Mediterr 7:70–71 Bertaccini A, Paltrinieri S, Contaldo N, Cavicchi L, Mori N, Bellardi MG (2015) Severe diseases induced by viruses and phytoplasmas in Hydrangea in Italy. Acta Hortic 1072:105–111 Brierley P (1954) Symptoms in the florist’s Hydrangea caused by Tomato ring spot virus and an unindentified sap transmissible virus. Phytopathology 44:696–699 Brierley P, Lorentz P (1957) Hydrangea ring spot virus, the probable cause of ‘running out’ of the florists’ hydrangea. Phytopathology 47:39–43 Caballero JEM, Lockhart BE, Mason SL, Daughtrey M (2009) Identification and properties of a carlavirus causing chlorotic mottle of florists’ hydrangea (H. macrophylla) in the United States. Plant Dis 93:891–895 Chiko AW, Godkin SE (1986) Occurrence of Alfalfa mosaic, Hydrangea ringspot and Tobacco ring spot viruses in Hydrangea spp. in British Columbia. Plant Dis 70:541–544 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among greenhouse ornamentals in Pennsylvania. Plant Dis 76:795–800 Hughes PL, Lawrence JE, Scott SW (2005) The complete nucleotide sequence of hydrangea ringspot virus. Arch Virol 150(11):2339–2345 Lawson RH (1995) Hydrangea. In: Loebenstein G, Lawson RH, Brunt AA (eds) Virus and virus-like diseases of bulb and flower crops. Wiley Publishers, UK, pp 484–492 Lockhart BEL, Mollov D, Daughtrey M (2013) First report of Alfalfa mosaic virus occurrence in hydrangea in the United States. Plant Dis 97(9):1258 Louro D (1996) Detection and identification of tomato spotted wilt virus and impatiens necrotic spot virus in Portugal. Acta Hortic 431:99–105 Mertelik J, Kloudova K (2009) First detection of Hydrangea ringspot virus in mountain hydrangea in the Czech Republic. Plant Pathol 58:405 Mertelik J, Kloudova K (2011) Hydrangea ring spot virus in Hydrangea spp. plants in the Czech Republic. Acta Hortic (ISHS) 901:237–238 Ruter JM, Gitaitis RD (1993) Impatiens necrotic spot virus in woody landscape plants in Georgia. Plant Dis 77:318 Song EG, Lee HS, Ryu KH (2016) Occurrence of hydrangea ringspot virus and hydrangea chlorotic mottle virus in hydrangea plants in South Korea. J Gen Plant Pathol 82:281–285

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Hylocereus spp. (Hylocereus undatus; H. polyrhizus) (Pitahaya)

Tamura M, Komuro Y (1967) The mosaic disease of hydrangea caused by cucumber mosaic virus. Ann Phytopathol Soc Jpn 33:27–31 Tang J, Harper SJ, Wei T, Clover GR (2010) Characterization of Hydrangea chlorotic mottle virus, a new member of the genus Carlavirus. Arch Virol 155(1):7–12 Thomas BJ, Barton RJ, Tuszynski A (1983) Hydrangea mosaic virus, a new illarvirus from Hydrangea macrophylla (Saxifragaceae). Ann Appl Biol 103:261–270 Veerakone S, Liefting LW, Lebas BSM, Ward L (2012) First report of Cherry leaf roll virus in Hydrangea macrophylla. Plant Dis 96:463 Welvaert W, Samyn G (1973) De verspreiding van het Hydrangea ring spot virus by de Hydrangea macrophylla (Thunb.) BC. Meded Faikulteir Landbouwwetensehappen Gent 38:1647–1654

Hylocereus spp. (Hylocereus undatus; H. polyrhizus) (Pitahaya) Family: Cactaceae

Cactus virus X Taxonomic position Genus: Potexvirus

Edible fruit

(CVX)

Family: Alphaflexiviridae

CVX infection in plants of Hylocereus spp. was reported from Taiwan, China, Korea, and Brazil (Hung et al. 1999; Liou et al. 2001, 2004; Liao et al. 2002; Duarte et al. 2008; Kim et al. 2016; Peng et al. 2016). The virus-infected pitahaya plants show chlorotic spots and mosaic symptoms. The virus is mechanically sap-transmissible, and the virus is also graft-transmissible (Duarte et al. 2008). For more details of CVX, refer to Opuntia spp.

Pitaya virus X Taxonomic position Genus: Potexvirus

(PiVX)

Family: Alphaflexiviridae

Geographical distribution PiVX infection in plants of Hylocereus spp. was reported from Taiwan (Mao 2008). Transmission The virus is transmitted by means not involving a vector. The virus is transmitted by mechanical sapinoculation. Use of infected planting material is the primary source of virus spread. Virion properties and genome Virions are flexuous filaments, 450–500 nm in length and 12–13 nm in diameter. The genome consists of a single molecule of linear positive-sense single-stranded RNA of 6677 nt (JF930327 = NC_024458)

Hylocereus spp. (Hylocereus undatus; H. polyrhizus) (Pitahaya)

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and comprises of five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunit is of one type, 18–27 kDa in size (Mao 2008; Li 2010).

Schlumbergera virus X Taxonomic position Genus: Potexvirus

(SchVX)

Family: Alphaflexiviridae

SchVX was reported from Brazil in plants of Hylocereus undatus showing mottling or chlorotic spotting (Duarte et al. 2008). The virus is transmissible by mechanical sap-inoculation, and also by grafting. For more details of SchVX, refer to Schlumbergera spp. (Syn.) Zygocactus spp.

H Zygocactus virus X Taxonomic position Genus: Potexvirus

(ZyVX)

Family: Alphaflexiviridae

ZyVX infection in plants of Hylocereus undatus was reported from Brazil and Taiwan (Duarte et al. 2008; Mao 2008). The virus infected pitahaya plants show mottling or chlorotic spotting. The virus is mechanically sap-transmissible and also by grafting. For more details of ZyVX, refer to Schlumbergera spp. (Syn.) Zygocactus spp.

References Duarte LML, Alexandre MAV, Rivas B, Harakava R, Galleti SR, Brradas MM (2008) Potexvirus diversity in Cactaceae from São Paulo State in Brazil. J Plant Pathol 90:545–551 Hung CL, Liou MR, Liou RF (1999) Identification and characterization of a Potexvirus on Pitaya (Hylocereus undatus). (Abs.). Plant Pathol Bull 8:177 Kim JS, Park CY, Nam M, Lee JS, Kim HG, Lee SH (2016) First report of Cactus virus X infecting Hylocereus polyrhizus in Korea. Plant Dis 100:2544 Li Y-S (2010) Characterization, infectious clone construction and antiserum preparation of Pitaya virus X. MS thesis. National Taiwan Univ. (In Chinese) Liao JY, Chang CA, Yan CR, Chen YC, Deng TC (2002) Detection and incidence of Cactus virus X on pitaya in Taiwan. Plant Pathol Bull 12:225–234 Liou MR, Hung CL, Liou RF (2001) First report of Cactus virus X on Hylocereus undatus (Cactaceae) in Taiwan. Plant Dis 85:229 Liou MR, Chen YR, Liou RF (2004) Complete nucleotide sequence and genome organization of a Cactus virus X strain from Hylocereus undatus (Cactaceae). Arch Virol 149:1037–1043 Mao CH (2008) Molecular characterization and detection of new Zygocactus virus X and Pitaya virus X from pitaya. MS thesis. National Taiwan Univ. (In Chinese) Peng C, Yu NT, Luo W, Fan HY, He F, Li XH, Zhang ZL, Liu ZX (2016) Molecular identification of Cactus virus X infecting Hylocereus polyrhizus (Cactaceae) in Hainan Island, China. Plant Dis 100:1956

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Hymenocallis littoralis (Beach spider lily)

Hymenocallis littoralis (Beach spider lily) Family: Amaryllidaceae

Ornamental

Calla lily chlorotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(CCSV)

Family: Tospoviridae

CCSV was reported infecting plants of Hymenocallis littoralis from China (Liu et al. 2012). The virusinfected beach spider lily plants exhibit systematic leaf necrosis, chlorotic lesions, rugosity, and deformation symptoms. The virus is mechanically transmissible to a variety of herbaceous hosts, but is naturally transmitted by a vector Thrips palmi in a persistent-propagative manner. For more details of CCSV, refer to Zantedeschia spp.

Hippeastrum chlorotic ringspot virus

(HCRV)

Taxonomic position HCRV is a tentative member of the genus Orthotospovirus and family Tospoviridae HCRV was reported in plants of Hymenocallis littoralis (spider lily) from China (Xu et al. 2013), causing concentric rings, leaf deformation, and chlorotic local lesions that developed into necrotic spots and mosaic. The virus is transmitted by thrips vectors, Thrips palmi and T. tabaci, in a persistentpropagative manner. For more details of HCRV, refer to Hippeastrum spp.

Hippeastrum mosaic virus Taxonomic position Genus: Potyvirus

(HiMV)

Family: Potyviridae

HiMV was reported infecting plants of Hymenocallis littoralis from the UK (Brunt 1973). The virus-infected beach spider lily plants exhibit symptoms of irregular mosaic in leaves and flowering stems. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of HiMV, refer to Hippeastrum spp.

Hyacinth mosaic virus Taxonomic position Genus: Potyvirus

(HyaMV)

Family: Potyviridae

HyaMV infection in plants of Hymenocallis littoralis was reported from the Netherlands (Pham et al. 2011). The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of HyaMV, refer to Hyacinth spp.

Hymenocallis littoralis (Beach spider lily)

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Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Hymenocallis littoralis was reported from Kunming, China (Liu et al. 2010). The virus-infected beach spider lily plants show chlorotic ringspots and necrotic spots. The virus is transmitted by thrips vectors in a persistent-propagative manner, and is also transmissible by mechanical inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Lily symptomless virus Taxonomic position Genus: Carlavirus

(LSV)

Family: Betaflexiviridae

LSV infection in plants of Hymenocallis littoralis (beach spider lily) was reported from India, showing symptoms of leaf curl, chlorotic striping, and chlorosis (Singh et al. 2005). The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of LSV, refer to Lilium spp.

Nerine yellow stripe virus Taxonomic position Genus: Potyvirus

(NeYSV)

Family: Potyviridae

An isolate of NeYSV infecting plants of Hymenocallis littoralis was reported and partially sequenced (EF362622) from the Netherlands (Pham et al. 2011). The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of NeYSV, refer to Nerine spp.

References Brunt AA (1973) Report Glasshouse Crops Research Institute for 1972, p 103 Liu YT, Zheng YX, Li YZ, Li ZY (2010) First report of impatiens necrotic spot virus on Spiderlily in China. Plant Dis 94:484 Liu Y, Lu X, Zhi L, Zheng Y, Chen X, Xu Y, Wu F, Li Y (2012) Calla lily chlorotic spot virus from spider lily (Hymenocallis litteralis) and tobacco (Nicotiana tabacum) in the South-west of China. J Phytopathol 160:201–205 Pham KTK, de Kock MJD, Lemmers MEC, Derks AFLM (2011) Molecular identification of potyviruses infecting bulbous ornamentals by the analysis of coat protein (CP) sequences. Acta Hortic 901:167–172 Singh AK, Hallan V, Verma N, Zaidi AA (2005) Natural occurrence of Lily symptomless virus on spider lily. Plant Pathol 54:255 Xu Y, Lou SG, Li XL, Zheng YX, Wang WC, Liu YT (2013) The complete S RNA and M RNA nucleotide sequences of a hippeastrum chlorotic ringspot virus (HCRV) isolate from Hymenocallis littoralis (Jacq.) Salisb in China. Arch Virol 158:2597–2601

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Hyoscyamus spp. (Hyoscyamus muticus, H. niger) (Henbane)

Hyoscyamus spp. (Hyoscyamus muticus, H. niger) (Henbane) Family: Solanaceae

Medicinal

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Hyoscyamus muticus was reported from India (Zaim and Khan 1988; Samad et al. 2000). The virus-infected henbane plants exhibit mottle crinkle and severe mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Henbane mosaic virus Taxonomic position Genus: Potyvirus

(HMV)

Family: Potyviridae

Geographical distribution HMV infection in plants of Hyoscyamus niger was reported from Egypt, Germany, Hungary, Italy, and the UK (Hamilton 1932; Watson 1937; Salamon and Palkovics unpublished - AM184113; El-Baz et al. unpublished - KM497011). Symptoms and host(s) The virus-infected henbane plants exhibit vein-clearing and green/yellow mosaic symptoms. The natural host of this virus in Hungary is Physalis alkekengi. Transmission The virus is transmitted by aphid vectors, Myzus persicae, M. ascalonicus, Aulacorthum circumflexum, and Macrosiphum euphorbiae, in a non-persistent manner. The virus is also transmissible by mechanical inoculation. Virion properties and genome The virions are non-enveloped, flexuous filaments, with no clear modal length of 800–900 nm and 12 nm wide. The genome consists of a single molecule of linear positive-sense single-stranded RNA of 9.5–10 kb. Partial genome sequences are available (KM497011; AM184113) (Revers and Garcia 2015; Wylie et al. 2017).

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References Hamilton MA (1932) On three new virus diseases of Hyoscyamus niger. Ann Appl Biol 19:550–567 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Samad A, Raj SK, Srivastava A, Chandra G, Ajayakumar PV, Zaim M, Singh BP (2000) Characterization of an Indian isolate of Cucumber mosaic virus infecting Egyptian henbane (Hyoscyamus muticus L.). Acta Virol 44(3):131–136 Watson MA (1937) Field experiments on the control of aphis-transmitted virus diseases of Hyoscyamus niger. Ann Appl Biol 24:557–573 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Zaim M, Khan MMAA (1988) Green mosaic of Hyoscyamus niger L. and Rauvolfia serpentina Benth. Indian J Plant Pathol 6:152–157

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I

Iberis semperflorens Family: Brassicaceae

Ornamental

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Iberis semperflorens was reported from Italy (Parrella et al. 2003, 2013). The virus-infected iberis plants exhibit symptoms of chlorotic/necrotic spots and rings. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Parrella G, Gognalons P, Gebre-Selassiè K, Vovlas C, Marchoux G (2003) An update on host-range of Tomato spotted wilt virus. J Plant Pathol 85:227–264 Parrella G, Cavicchi L, Greco B, Bellardi MG (2013) First report of Tomato spotted wilt virus in Iberis semperflorens. J Plant Pathol 95:S4.74

Ibicella lutea (Devil’s claw) Synonyms Martynia lutea; Proboscidea lutea Family: Martyniaceae

Ornamental

© Springer Nature India Private Limited 2019 K. S. Sastry et al., Encyclopedia of Plant Viruses and Viroids, https://doi.org/10.1007/978-81-322-3912-3

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Ilex crenata (Japanese holly)

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Ibicella lutea was reported from Iran (Safaeizadeh and Saidi 2012). The virus-infected devil’s claw plants exhibit mosaic, mottling, and leaf malformation symptoms. The virus is transmitted by a number of aphid vectors in a non-persistent manner, and also by mechanical sapinoculation to a large number of host plants. For more details of CMV, refer to Cucumis sativus.

References Safaeizadeh M, Saidi A (2012) First report of Cucumber mosaic virus on Ibicella lutea in Iran. J Plant Pathol 94:S4.95

Ilex crenata (Japanese holly) Family: Aquifoliaceae

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

Ornamental

(TRSV)

Family: Secoviridae

TRSV infection in plants of Ilex crenata was reported from Maryland, USA (Waterworth and Povish 1977). The virus-infected Japanese holly plants exhibit symptoms of chlorotic yellow spots in young leaves and irregular margins in older leaves, and leaves are permanently distorted although plant growth was not appreciably reduced. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

References Waterworth HE, Povish WR (1977) A yellow leaf spot disease of Ilex crenata caused by Tobacco ring spot virus. Plant Dis Reptr 61:104–105

Ilex glabra (Appalachian tea, Inkberry) Family: Aquifoliaceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

(INSV)

Ilex paraguariensis (Yerba mate)

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INSV infection in plants of Ilex glabra was reported from the Georgia, USA (Ruter and Gitaitis 1993). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

References Ruter JM, Gitaitis RD (1993) Impatiens necrotic spot virus in woody landscape plants in Georgia. Plant Dis 77:318

Ilex paraguariensis (Yerba mate) Family: Aquifoliaceae

Commercial crop

Yerba mate alphaendornavirus 1 Taxonomic position Genus: Alphaendornavirus

(YmEV 1)

Family: Endornaviridae

Geographical distribution YmEV 1 infection in plants of Ilex paraguariensis was reported from Argentina (Debat et al. 2014). Symptoms and host(s) The virus-infected yerba mate plants do not exhibit visible symptoms. Transmission The virus is transmitted through the seed via both the ova and pollen. No horizontal spread has been observed in the field and no potential vectors have been identified. The virus is not mechanically saptransmissible. Virion properties and genome The virus does not have a gene for capsid protein; therefore no virion is formed. The complete sequence of the linear dsRNA viral genome is 13,954 bp, consists of a single 13,743 bp ORF, with a 149 bp 50 UTR and a 61 bp 30 UTR (KJ634409). The predicted ORF encodes a 4581 aa polypeptide with a UDPglucose glycosyl transferase, a capsular polysaccharide synthesis protein, and a RNA-dependent RNA polymerase domain. The dsRNA genome is encapsulated along with the viral replicase enzyme. A sitespecific break (nick) is found in the coding strand about 1–2 kb from the 50 terminus.

Yerba mate chlorosis-associated virus

(YmCaV)

Taxonomic position YmCaV is a tentative member of the genus Cytorhabdovirus and family Rhabdoviridae Geographical distribution YmCaV infection in plants of Ilex paraguariensis was reported from Argentina (Bejerman et al. 2017).

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Impatiens spp.

Transmission The virus is transmitted by leafhopper vectors. Virion properties and genome The virions are bullet-shaped. The genome is monopartite, negative-sense, single-stranded RNA of 12876 nt (KY366322) (Walker et al. 2018).

References Bejerman N, de Breuil S, Debat H, Miretti M, Badaracco A, Nome C (2017) Molecular characterization of yerba mate chlorosis-associated virus, a putative cytorhabdovirus infecting yerba mate (Ilex paraguariensis). Arch Virol 2017 162(8):2481–2484 Debat HJ, Grabiele M, Aguilera PM, Bubillo R, Zapata PD, Marti DA, Ducasse DA (2014) The complete genome of a putative endornavirus identified in yerba mate (Ilex paraguariensis St. Hil.). Virus Genes 49:348–350 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Rhabdoviridae. J Gen Virol 99:447–448

Impatiens spp. Family: Balsaminaceae

Ornamental

Citrus exocortis viroid

(CEVd)

Taxonomic position Genus: Pospiviroid

Family: Pospiviroidae

CEVd infection in plants of Impatiens spp. was reported from Canada and India (Nie et al. 2005; Singh and Baranwal 2006; Singh et al. 2009). The viroid is transmissible through the infected vegetative propagative material. A high seed transmission rate was observed in case of CEVd in impatiens (Singh et al. 2009). The presence of CEVd in non-germinated seed was 21%, while the transmission rate in seedlings was 66% in Impatiens walleriana. Following 2 years of seed storage, the respective figures were 6% and 26% (Baranwal et al. 2009; Singh et al. 2009). For more details of CEVd, refer to Citrus spp.

Clover yellow vein virus Taxonomic position Genus: Potyvirus

(ClYVV)

Family: Potyviridae

ClYVV infection in plants of Impatiens sultani was reported from Japan (Chun et al. 1990; Xiang et al. 1990). The virus-infected impatiens plants exhibit foliar necrotic ring symptoms. The virus is transmitted by several aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ClYVV, refer to Trifolium spp.

Impatiens spp.

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Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Impatiens spp. was reported from South Carolina (USA), and Poland (Herold 1964; Polak 1967; Flasinski et al. 1995; Kaminska 1995). The virus-infected impatiens plants exhibited symptoms of mosaic and deformation of leaves, and plants were stunted. Some impatiens cultivars infected with CMV exhibited leaf narrowing, with ragged leaf borders, as well as split petals and color breaking. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Helenium virus S

(HVS)

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Synonyms Impatiens latent virus Taxonomic position Genus: Carlavirus

Family: Betaflexiviridae

HVS was reported in plants of Impatiens holstii (syn. I. walleriana) in the USA (initially as Impatiens latent virus; Lockhart and Betzold 1980a; Chun et al. 1990) and Europe, causing unobtrusive symptoms. Although the helenium isolate was shown to be aphid-transmitted, no aphid transmission was obtained with the impatiens isolate (Koenig et al. 1983; Plese et al. 1988). The virus is mechanically sap-transmissible. For more details of HVS, refer to Helenium spp.

Impatiens flower break virus Taxonomic position Genus: Potyvirus

(IFBV)

Family: Potyviridae

Geographical distribution IFBV infection in plants of Impatiens spp. was reported from the USA (Jordan and Guaragna 2007; Jordan et al. 2011; Hammond et al. 2011) and Korea (Cho et al. 2017). Symptoms and host(s) The virus-infected impatiens plants exhibit colour breaking symptoms on flower petals. Transmission The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation.

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Impatiens spp.

Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long and 11–13 nm wide. The genome contains single-molecule of linear, positive-sense single-stranded RNA of 9622 nt in length (KU981084 = NC_030236) (Wylie et al. 2017).

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

Geographical distribution INSV was first reported in plants of New Guinea impatiens (Impatiens spp.) in the USA (Law and Moyer 1990). The virus also occurs in Canada, Mexico, Germany, the Netherlands, Poland, Belgium, Iran, the Czech Republic, Hungary, Israel, the UK, and also in many Southeast Asian countries (Mertelik et al. 2002; Shahraeen et al. 2002; Lebas et al. 2004; CABI/EPPO 2007; Toth et al. 2007; Ghotbi 2013). Symptoms and host(s) Characteristic symptoms of INSVon naturally infected seed-type impatiens include small black to brown necrotic flecks and ring-spots. Small dark purple ring-spots occur on leaves and white rings may be present on flower petals. Necrosis may also appear along the midribs of diseased leaves that often show yellowing, malformation, and stunting. Symptoms of necrosis may also appear in the stems of affected plants. Infected leaves often drop prematurely. Temperature and stage of the growth of the host have a significant effect on symptom expression of INSV (Law and Moyer 1990; Daughtrey et al. 1995). A few of the many hosts of INSV include impatiens, begonia, vinca, cineraria, exacum, cyclamen, chrysanthemum, and alstroemeria; but there are many more flowering plants, vegetables, and weeds that can host this virus (Daughtrey et al. 1997). Transmission The virus is transmitted by the thrips vectors, Frankliniella occidentalis, F. intonsa, and F. fusca in a persistent-propagative manner (DeAngelis et al. 1993; Naidu et al. 2001; Sakurai et al. 2004). The virus multiplies in adult thrips, which can transmit the virus within 2–3 days after feeding on infected plants, giving infection rates of up to 95% (Wijkamp and Peters 1993). Virus transmission and movement from plant to plant occur due to feeding activity of the thrips. Larvae acquire the virus by feeding on an infected host plant. The virus is mechanically sap-transmissible to more than 648 different plant species (Daughtrey et al. 1997). Nicotiana benthamiana and Petunia hybrida are used as bioassay hosts to detect the virus. Necrotic spot local lesions are formed on the leaves of Nicotiana benthamiana followed by systemic vein yellowing and necrosis. Petunia leaves show necrotic lesions on inoculated leaves without systemic symptoms. The virus infects predominantly ornamentals such as begonia, carnation, chrysanthemum, dahlia, geranium, gladiolus, impatiens, narcissus, orchids, petunia, etc. Virion properties and genome The virions are spherical and membrane bound measuring about 80–120 nm in diameter. The genome is comprised of three unique molecules of negative or ambisense ssRNA, designated L (large) 8776 nt (X93218 = NC_003625), M (medium) 4972 nt (M74904 = NC_003616), and S (small) 2992 nt (X66972 = NC_003624) (de Haan et al. 1992; Law et al. 1992; van Poelwijk et al. 1997).

Impatiens spp.

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Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

IYSV infection in plants of Impatiens spp. was reported from the Netherlands (Gera et al. 2008). The virus-infected impatiens plants exhibit symptoms of necrotic rings on leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of IYSV, refer to Iris spp. or Allium cepa.

Ligustrum necrotic ringspot virus Taxonomic position Genus: Carlavirus

(LNRSV)

Family: Betaflexiviridae

LNRSV was reported from plants of Impatiens spp. in New Zealand (GQ411367) and the USA (Harper et al. unpublished - GQ411367; Jordan unpublished). The virus is mechanically sap-transmissible and also transmissible by grafting. For more details of LNRSV, refer to Ligustrum spp.

Ribgrass mosaic virus Taxonomic position Genus: Tobamovirus

(RMV)

Family: Virgaviridae

RMV infection in plants of New Guinea Impatiens was reported from Germany (Wetzel et al. 2006). The virus-infected impatiens plants exhibit severe leaf chlorotic mottling and deformation symptoms. No vector is reported for this virus. The virus is mechanically sap-transmissible. The virus is transmissible by grafting and by contact between plants. For more details of RMV, refer to Plantago lanceolata.

Strawberry latent ringspot virus Taxonomic position Genus: Unassigned

(SLRSV)

Family: Secoviridae

SLRSV infection in plants of Impatiens walleriana was reported from New Zealand (Tang et al. 2013). The virus-infected impatiens plants exhibit symptoms of chlorotic streaks and necrotic rings on the leaves. The virus is transmitted by nematode vectors (Xiphinema spp.) and also by mechanical sapinoculation. The virus is transmissible by grafting. For more details of SLRSV, refer to Fragaria spp.

Tobacco mild green mosaic virus Taxonomic position Genus: Tobamovirus

(TMGMV)

Family: Virgaviridae

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Impatiens spp.

TMGMV was reported in plants of Impatiens spp. from the UK (Skelton et al. 2010; Harju et al. 2011). The virus-infected impatiens plants exhibited symptoms of leaf necrosis, distorted leaves, foliar reddening on older plants, necrotic lesions, and paler flower color; the plants were also stunted. Pinkor orange-flowering varieties exhibit flecking of flowers. No insect vector is reported for this virus. The virus is mechanically sap-transmissible and has a wide host range. The virus is transmissible by grafting and also by contact between plants. For more details of TMGMV, refer to Nicotiana tabacum.

Tobacco mosaic virus

(TMV)

Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

TMV infection in plants of Impatiens hawkeri was reported from Brazil (Rivas et al. 2000). The virusinfected impatiens plants exhibit chlorotic spots and rings and leaf deformation symptoms. There is no known vector for this virus. The virus is mechanically sap-transmissible and also transmissible by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Impatiens holstii was reported from Minnesota and Iowa (USA), and from Iran (Lockhart and Pfleger 1979; Ghotbi and Shahraeen 2012). The virus-infected Impatiens holstii plants showed stunting leaf mottling and premature leaf abscission. Virus-infected Impatiens walleriana plants show symptoms of stunting, mosaic, chlorotic ring-spots, and line patterns. In some cultivars, systemic symptoms of mosaic and ring-spots were observed. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Impatiens holstii was reported from Minnesota, USA (Lockhart and Betzold 1980b). The virus-infected impatiens plants show symptoms of leaf twisting, deformation, and stunting. Flowers from infected plants have not shown any symptoms. The virus is transmitted by thrips vectors; the virus present in/on pollen entering the host through injuries caused by thrips while feeding. The virus is mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Impatiens spp.

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Tomato mosaic virus

(ToMV)

Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

ToMV infection in plants of Impatiens hawkeri was reported from Brazil (Duarte et al. 2003, 2007). The virus-infected impatiens plants exhibit mosaic and leaf distortion symptoms. No vector is involved in the spread of this virus. This virus is transmissible by mechanical inoculation, by grafting, and by contact between plants. For more details of ToMV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV occurs in plants of Impatiens spp. in the Netherlands, Serbia and elsewhere in Europe, Canada, and the USA (Matteoni et al. 1988; Hsu and Lawson 1991; De Avila et al. 1992; German et al. 1992; Hausbeck et al. 1992; Daughtrey et al. 1997; Dekic et al. 2008). The virus-infected impatiens plants exhibit small, dark purple-brown, yellow, or brown spots or rings and also brown lines, streaks, or zigzag patterns on the leaves. Flowers may also show white ring-spots. Dark streaks appear on stems and leaves, especially on or near the midrib. Leaf yellowing, leaf distortion, leaf drop, and stunting may occur. As the virus becomes systemic, faint purplish ring patterns develop on the lower leaves. Young infected impatiens plants, or older affected plants, may drop many of their leaves, appear “leggy” and are generally unthrifty. Some cultivars of New Guinea impatiens die; others exhibit uneven growth which reduces their value. The virus is mechanically sap-transmissible to a large number of hosts and is transmitted by thrips vectors, viz., Frankliniella schulzei, F. fusca, and Thrips tabaci, in a persistentpropagative manner (Wijkamp et al. 1996). For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow ring virus

(TYRV)

Taxonomic position TYRV is a tentative member of the genus Orthotospovirus and family Tospoviridae TYRV was reported in plants of Impatiens spp. in Iran (Ghotbi and Shahraeen 2012). The virus is transmitted by a thrips vector, Thrips tabaci, in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of TYRV, refer to Solanum lycopersicum.

Turnip mosaic virus

(TuMV)

Synonyms Cabbage black ringspot virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

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Impatiens spp.

TuMV infection in plants of Impatiens balsamina was reported from Geneva (Switzerland) and Slovakia (Polak 1967; Provvidenti 1982; Farzadfar et al. 2005). The virus-infected I. balsamina plants showed black stem streaks and foliar chlorosis with necrotic veins and spots. Few plants reached full maturity, but the majority of infected plants collapsed and died prematurely. Some varieties of impatiens ‘New Guinea hybrids’ exhibit dark-colored areas at the base or tip of the leaves, and leaves may also be asymmetric. The virus is transmitted by a large number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

References Baranwal VK, Singh RP, Dilworth AD, Ao D, Xiaoping, Singh M (2009) Citrus exocortis viroid transmission through commercially – distributed seeds of Impatiens and Verbena plants. Eur J Plant Pathol 124:691–694 CABI/EPPO (2007) Impatiens necrotic spot virus. Distribution maps of plant diseases, no. 755. CAB International, Wallingford Cho S-Y, Kim H, Yi S-I, Lim S, Park JM, Cho HS, Kim H, Kwon S-Y, Moon JS (2017) First report of Impatiens flower break virus infecting Impatiens walleriana in South Korea. Plant Dis 101:394 Chun XB, Ohki ST, Osaki T, Inouye T (1990) Clover yellow vein virus and a carlavirus isolated from Impatiens sultani in Japan. Ann Phytopathol Soc Jpn 56:557–560 Daughtrey ML, Wick RL, Peterson JL (1995) Compendium of flowering potted plant diseases. APS Press, St. Paul Daughtrey M, Jones RK, Moyer JW, Daub ME, Baker JR (1997) Tospoviruses strike the greenhouse industry – INSV has become a major pathogen on flower crops. Plant Dis 81:1220–1230 DeAngelis JD, Sether DM, Rossignol PA (1993) Survival, development and reproduction in western flower thrips (Thysanptera: Thripidae) exposed to Impatiens necrotic spot virus. Environ Entomol 22:1308–1312 De Avila AC, De Haan P, Kitajima EW, Kormelink R, Resende R, Goldbach RW, Peters D (1992) Characterization of a distinct isolate of Tomato spotted wilt virus (TSWV) from Impatiens sp. in the Netherlands. J Phytopathol 134:133–151 de Haan P, de Avila AC, Kormelink R, Westerbroek A, Gielen JJ, Peters D, Goldbach R (1992) The nucleotide sequence of the S RNA of Impatiens necrotic spot virus, a novel tospovirus. FEBS Lett 306(1):27–32 Dekic I, Bulajic A, Vucurovic A, Ristic D, Krstic B (2008) Influence of Tomato spotted wilt virus uneven distribution on its serological detection in tomato, pepper and ornamentals. Pestic Phytomed 23:225–234 Duarte LML, Soares RM, Fernandes FMC, Alexandre MAV, Tozetto ARP, Cilli A, Rivas EB (2003) Tomato mosaic virus isolado de Impatiens hawkeri: análise filogenética molecular, vol 70. Arquivos do Instituto Biológico, São Paulo, pp 122–127 Duarte LML, Alexandre MAV, Rivas EB, Cattai MB, Soares RM, Harakava R, Fernandes FMC (2007) Phylogenetic analysis of Tomato mosaic virus from Hemerocallis sp. and Impatiens hawkeri. Summa Phytopathol 33:409–413 Farzadfar SH, Ohshima K, Pourrahim R, Golnaraghi AR, Jalali S, Ahoonmanesh A (2005) Occurrence of Turnip mosaic virus on ornamental crops in Iran. Plant Pathol 54:261 Flasinski S, Scott SW, Barnett OW, Sun C (1995) Diseases of Peperomia, Impatiens, and Hibbertia caused by Cucumber mosaic virus. Plant Dis 79:843–848 Gera A, Siti L, Beckelman Y, Tam Y, Kritzman A, Zeidan M (2008) First report of Iris yellow spot tospovirus (IYSV) in lily and impatiens. In: Proceedings of the 12th international symposium on virus diseases of ornamental plants, Haarlem, 2008-04-20/24, p 51 German TL, Ullman DE, Moyer JW (1992) Tospoviruses: diagnosis, molecular biology, phylogeny, and vector relationships. Ann Rev Phytopathol 30:315–348 Ghotbi T (2013) Serological and molecular detection of INSVand introduction of some INSVornamental host plants from five provinces in Iran. Iran J Plant Pathol 49:41 Ghotbi T, Shahraeen N (2012) Incidence and distribution of viruses infecting propagated ornamentals in Northern Iran. Int Res J Microbiol 3:373–381 Hammond J, Jordan R, Vaira AM (2011) Virus diseases of ornamentals, in soils, plant growth and crop production. In: UNESCO-EOLSS Joint Committee (ed) Encyclopedia of Life Support Systems (EOLSS), Developed under the auspices of the UNESCO. Eolss Publishers, Oxford. http://www.eolss.net Harju VA, Skelton AL, Monger WA, Forde SMD, Daly M, Nixon T, Lawson R, Bennett S, Mumford RA (2011) A report of viruses, viroids and phytoplasmas found in ornamental plants from 1999–2007: findings from central science laboratory, UK. Acta Hortic 901:223–229

Impatiens spp.

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Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among green house ornamentals in Pennsylvania. Plant Dis 76:795–800 Herold F (1964) Natural infection of Impatiens sultani with Cucumber mosaic virus. Plant Dis Reptr 48:603–605 Hsu HT, Lawson RH (1991) Direct tissue blotting for detection of tomato spotted wilt virus in impatiens. Plant Dis 75:292–295 Jordan R, Guaragna M (2007) Characterization of a new Potyvirus, Impatiens flower break virus, infecting New Guinea Impatiens. Phytopathology 97:S54 Jordan R, Guaragna MA, Putnam M (2011) Detection and molecular characterization of new and emerging potyviruses of ornamental plants. Acta Hortic 901:159–166 Kaminska M (1995) Natural occurrence of cucumber mosaic virus in Impatiens sp. ‘New Guinea’. Phytopathol Pol 22:21–27 Koenig R, Lesemann D-E, Lockhart B, Betzold JA, Weidemann HL (1983) Natural occurrence of Helenium virus S in Impatiens holstii. J Phytopathol 106:133–140 Law MD, Moyer JW (1990) A tomato spotted wilt-like virus with a serologically distinct N protein. J Gen Virol 71:933–938 Law MD, Speck J, Moyer JW (1992) The M RNA of Impatiens necrotic spot tospovirus (Bunyaviridae) has an ambisense genomic organization. Virology 188:732–741 Lebas BSM, Ochoa-Corona FM, Elliott DR, Tang Z, Alexander BJR, Froud KJ (2004) An investigation of an outbreak of Impatiens necrotic spot virus in New Zealand. Phytopathology 94:S57 Lockhart BEL, Betzold JA (1980a) Latent infection of Impatiens by a potexvirus. Acta Hortic 110:81–84 Lockhart BEL, Betzold JA (1980b) Leaf curl of impatiens caused by Tobacco streak virus infection. Plant Dis 64:289–290 Lockhart BEL, Pfleger FL (1979) Identification of a strain of Tobacco ring spot virus causing a disease of impatiens in commercial greenhouses. Plant Dis Reptr 63:258–261 Matteoni JA, Allen WR, Broadbent AB (1988) Tomato spotted wilt virus in greenhouse crops in Ontario. Plant Dis 72:801 Mertelik J, Mokra V, Gotzova B, Gabrielova S (2002) Occurrence and identification of Impatiens necrotic spot tospovirus in the Czech Republic. Acta Hortic 568:79–83 Naidu RA, Deom CM, Sherwood JL (2001) First report of Frankliniella fusca as a vector of Impatiens necrotic spot tospovirus. Plant Dis 85:1211–1212 Nie X, Singh RP, Bostan H (2005) Molecular cloning, secondary structure, and phylogeny of three pospiviroids from ornamental plants. Can J Plant Pathol 27:592–602 Plese N, Erić Z, Krajačić M (1988) Further information on infection of Impatiens holstii with Helenium virus S. Acta Hortic 234:477–484 Polak Z (1967) Impatiens parviflora DC. a natural host of cabbage black ringspot and cucumber mosaic viruses. Biol Plant 9:354–359 Provvidenti R (1982) A destructive disease of garden balsam caused by a strain of Turnip mosaic virus. Plant Dis 66(11):1076–1077 Rivas EB, Pezani EF, Alexandre MAV, Duarte LML (2000) First report of a Tobamovirus in Dieffenbachia and Impatiens. Plant Dis 84:707 Sakurai T, Inoue T, Tsuda S (2004) Distinct efficiencies of Impatiens necrotic spot virus transmission by five thrips vector species (Thysanoptera: Thripidae) of tospoviruses in Japan. Appl Entomol Zool 39:71–78 Shahraeen N, Ghotbi T, Mehraban AH (2002) Occurrence of Impatiens necrotic spot virus in ornamentals in Mahallat and Tehran provinces in Iran. Plant Dis 86:694 Singh RP, Baranwal VK (2006) Symptomless expression of citrus exocortis viroid in ornamental plant species, Impatiens and Verbena, may be a potential threat to potato and tomato crops. Can J Plant Pathol 28:364–364 Singh RP, Dilworth AD, Xiaoping AO, Singh M, Baranwal VK (2009) Citrus exocortis viroid transmission through commercially – distributed seeds of Impatiens and Verbena plants. Eur J Plant Pathol 124:691–694 Skelton A, Nixon T, Monger W, Bennett S, Daly M, Hobden E, Harju V (2010) Tobacco mild green mosaic virus in Impatiens and Osteospermum: new hosts and first report in the UK. Plant Pathol 59:1160 Tang J, Ward LI, Clover GRG (2013) The diversity of Strawberry latent ringspot virus in New Zealand. Plant Dis 97:662–667 Toth EK, Kriston E, Takacs A, Bajtek M, Kazinczi G, Horvath J (2007) First report of Impatiens necrotic spot virus in ornamental plants in Hungary. Plant Dis 91:331 van Poelwijk F, Prins M, Goldbach R (1997) Completion of the impatiens necrotic spot virus genome sequence and genetic comparison of the L proteins within the family Bunyaviridae. J Gen Virol 78(Pt 3):543–546 Wetzel T, Njapo Ngangom HO, Chotewutmontri S, Krezal G (2006) Nucleotide sequence of a new isolate of Ribgrass mosaic tobamovirus infecting Impatiens New Guienea. Arch Virol 151:787–791 Wijkamp I, Peters D (1993) Determination of the median latent period of two Tospoviruses in Frankliniella occidentalis, using a novel leaf disk assay. Phytopathology 83:986–991 Wijkamp I, Van de Wetering F, Goldbach R, Peters D (1996) Transmission of Tomato spotted wilt virus by Frankliniella occidentalis: median acquisition and inoculation access period. Ann Appl Biol 129:303–313

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Imperata cylindrica (Blady grass)

Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Xiang BC, Ohki ST, Osaki T, Inouye T (1990) Clover yellow vein virus and a carlavirus isolated from Impatiens sultani in Japan. Ann Phytopathol Soc Jpn 56:557–560

Imperata cylindrica (Blady grass) Family: Poaceae

Weed host

Imperata yellow mottle virus Taxonomic position Genus: Sobemovirus

(IYMV)

Family: Solemoviridae

Geographical distribution IYMV infection in plants of Imperata cylindrica was reported from Burkina Faso, West Africa, by Tamm and Truve (2000) and Sereme et al. (2008). Symptoms and host(s) The virus-infected blady grass plants show yellow mottle symptoms; apart from I. cylindrica, the host range is restricted to Zea mays and experimentally to rice, sorghum, pearl millet, and the wild grasses Setaria verticillata and Rottboellia exaltata (Koala et al. 2017). Transmission The virus is transmissible by mechanical sap-inoculation; no vector has been identified, but beetle transmission is likely, as for other sobemoviruses. Virion properties and genome The virions are isometric, non-enveloped, and 32 nm in diameter. The virions contain a single coat protein of 29.1 kDa. The genome is a polycistronic positive-sense single-stranded RNA (ssRNA) of 4,547 nt (AM990928 = NC_011536). The genome consists of two overlapping ORFs, ORF2a and ORF2b (Sereme et al. 2008; Somera et al. 2015).

References Koala M, Traoré VSE, Sérémé D, Neya BJ, Brugidou C, Barro N, Traoré O (2017) Imperata yellow mottle virus: an emerging threat to maize, sorghum and pearl millet in Burkina Faso. Agric Sci 8(5):397–408 Sereme D, Lacombe S, Konate M, Pinel-Galzi A, Traore VSE, Hebrard E, Traore O, Brugidou C, Fargette D, Konate K (2008) Biological and molecular characterization of a new putative sobemovirus infecting Imperata cylindrica and maize in Africa. Arch Virol 153:1813–1820 Somera M, Sarmiento C, Truve E (2015) Overview on Sobemoviruses and a proposal for the creation of the family Sobemoviridae. Viruses 7:3076–3115 Tamm T, Truve E (2000) Sobemoviruses. J Virol 74:6231–6241

Inula spp. (Inula helenium, I. viscosa)

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Indigofera hirsuta (Hairy indigo) Family: Fabaceae

Weed host

Horsegram yellow mosaic virus Taxonomic position Genus: Begomovirus

(HgYMV)

Family: Geminiviridae

HgYMV infection in plants of Indigofera hirsuta was reported from India (Muniyappa and Reddy 1979). The virus-infected hairy indigo plants exhibit yellow mosaic symptoms on leaves and reduction of leaf size. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative nonpropagative manner. The virus is not transmissible by mechanical sap-inoculation, and not transmissible by contact between plants. For more details of HgYMV, refer to Macrotyloma uniflorum.

References Muniyappa V, Reddy HR (1979) Indigofera hirsutum a natural reservoir of Horsegram yellow mosaic virus. Madras Agric J 66:350

Inula spp. (Inula helenium, I. viscosa) Family: Asteraceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Medicinal plant

(CMV)

Family: Bromoviridae

CMV infection in plants of Inula viscosa was reported from Italy (EU432181) (Davino et al. 2010, 2012). The virus-infected inula plants exhibit symptoms of chlorotic mottling of the leaves. The virus is transmitted by a number of aphid vectors in a non-persistent manner, and also by mechanical sapinoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

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TSWV infection in plants of Inula helenium was reported from Bulgaria (Dikova 2011). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Davino S, Panno S, Davino M, Bellardi MG (2010) Inula viscosa new host of Cucumber mosaic virus. J Plant Pathol 92: S4.112 Davino S, Panno S, Rangel EA, Davino M, Bellardi MG, Rubio L (2012) Population genetics of Cucumber mosaic virus infecting medicinal, aromatic and ornamental plants from northern Italy. Arch Virol 157:739–745 Dikova B (2011) Tomato spotted wilt virus on some medicinal and essential oil-bearing plants in Bulgaria. Bulg J Agric Sci 17:306–313

Ipomoea batatas (Sweet potato) Family: Convolvulaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Tuber crop

(CMV)

Family: Bromoviridae

CMV occurs in Ipomoea batatas worldwide and is reported from China, Israel, Spain, Egypt, Japan, South Africa, New Zealand, Ghana, and West Africa (Martin 1962; Clark and Moyer 1988; Cohen and Loebenstein 1991; Opiyo et al. 2010; Qiao et al. 2012; Sossah et al. 2015). The virus-infected sweet potato plants exhibit symptoms include stunting, chlorosis, and yellowing. The virus is transmitted by a large number of aphids in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. Vegetative propagative material from infected plants plays a major role in virus spread (Cohen et al. 1988). For more details of CMV, refer to Cucumis sativus.

Sweet potato C3 virus

(SPC3V)

Taxonomic position SPC3V is a tentative member of the genus Phlebovirus and family Peribunyaviridae. Geographical distribution SPC3V was first reported in plants of Ipomoea batatas from Brazil and Uganda (Fuentes and Salazar 1989; Clark et al. 2012; Wassawa 2012). Symptoms and host(s) The virus-infected sweet potato leaves show deformation, vein-clearing, and mosaic symptoms. Symptoms vary from cultivar, viz., infected sweet potato cv. Paramo N and Gyno, show mottle symptoms and sweet potato cv. Georgia Red expressed red interveinal mottling.

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Transmission The virus failed to be transmissible through mechanical sap-inoculation, and also aphid Myzus persicae. The virus is graft-transmissible. The insect vector has not been identified. Virion properties and genome Flexuous virus particles are associated with this disease. Genome sequence information is not available.

Sweet potato C6 virus Taxonomic position Genus: Carlavirus

(SPC6V)

Family: Betaflexiviridae

Geographical distribution SPC6V infection in plants of Ipomoea batatas was reported from Dominican Republic, Peru, Kenya, South Africa, New Zealand, Uganda, Cuba, the Philippines, Indonesia, Egypt, Puerto Rico, and the USA (Fuentes 1994; Wassawa 2012). Symptoms and host(s) The virus-infected sweet potato plants exhibit chlorotic spots on the leaves and in some Ipomoea species exhibit chlorotic spotting and vein-clearing, mottling symptoms, and leaf dropping in the later stages. The natural host range of this virus is restricted to the Convolvulaceae. Transmission The virus is mechanically sap-transmissible at low efficiency and is also graft-transmissible. Virion properties and genome The virions are flexuous filaments about 610–700 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8857 nt (NC_018448, JX212747) and comprises 6 ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Adams et al. 2004; De Souza et al. 2013).

Sweet potato chlorotic fleck virus Taxonomic position Genus: Carlavirus

(SPCFV)

Family: Betaflexiviridae

Geographical distribution SPCFV infection in plants of Ipomoea batatas was reported from Peru, Kenya, New Zealand, China, Korea, Cuba, Panama, Bolivia, Brazil, Columbia, Cuba, Indonesia, Japan, Kenya, the Philippines, Uganda, Australia, Asia, Africa, South America, Ghana, and Austria (Usugi et al. 1991; Fuentes and Salazar 1992; Gibson et al. 1997; Makeshkumar et al. 2001; Mukasa et al. 2003a; Ateka et al. 2004a; Tairo et al. 2004; Jones and Dwyer 2007; Nyaboga et al. 2008; Kathurima et al. 2011; Qiao et al. 2012; Deng et al. 2014; Kwak et al. 2014; Sossah et al. 2015).

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Symptoms and host(s) The virus-infected sweet potato plants exhibit symptoms of chlorosis, leaf distortion, and vein-clearing. Transmission The virus is mechanically sap-transmissible to Convolvulaceae and Chenopodiaceae species. No seed transmission is observed (Aritua et al. 2009). Virion properties and genome The virions are flexuous filaments about 750–800 nm in length and 12 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 9104 nt (AY461421 = NC_006550) and comprises 6 ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Fuentes and Salazar 1992; Adams et al. 2004; Aritua et al. 2007b, 2009).

Sweet potato chlorotic stunt virus

(SPCSV)

Synonyms Sweet potato sunken vein virus Taxonomic position Genus: Crinivirus

Family: Closteroviridae

Geographical distribution SPCSV infection in Ipomoea batatas plants was first reported from Israel by Cohen et al. (1992). The virus occurs worldwide, being particularly prevalent in sub-Saharan Africa but with recent reports of damage in South America, Central America, Peru, Kenya, Ghana, Uganda, Tanzania, Egypt, Israel, Rwanda, China, and the USA. SPCSV has been differentiated into two distantly related strains, East African and West African, based on serology and nucleotide sequence data (Brown et al. 1988; Ngeve 1990; Wambugu 1991; Hoyer et al. 1996a, b; Milgram et al. 1996; Vetten et al. 1996; Gibson et al. 1998; Gibson and Aritua 2002; Yun et al. 2002; Ateka et al. 2004a; Lozano et al. 2004; Valverde and Moreira 2004; Valverde et al. 2004a; Abad et al. 2007; Njeru et al. 2008; Opiyo et al. 2010; Kathurima et al. 2011; Qiao et al. 2011, 2012; Hegde et al. 2012; Qin et al. 2013a; Sossah et al. 2015). Symptoms and host(s) The characteristic symptoms on sweet potato plants are stunting, chlorosis, and purpling of leaves. In addition to infecting sweet potato, SPCSV has also been transmissible to a range of Ipomoea spp. including I. setosa, I. acuminata, I. hederacea, I. hederifolia, I. nil cv. Scarlet O’Hara, I. purpurea, I. trichocarpa, I. trifida, I. wrightii, I. mexicana, I. alba (= I. bona-nox), and I. hildebrandtii. It has also been transmissible to Nicotiana clevelandii, N. benthamiana, Amaranthus palmeri, and to Lisianthus (Eustoma grandiflorum). Natural infection of SPCSV (reported by the name: SPSVV) was detected in Lisianthus (Eustoma grandiflorum) in the northern Negev in Israel (Cohen et al. 2001). In Lisianthus plants no leaf symptoms of SPSVV were observed, but flower stems were shorter by about one third. These data indicate that Lisianthus is a host for SPSVV.

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Transmission The virus is transmitted by whitefly vectors Bemisia tabaci, B. afer, and Trialeurodes vaporariorum in a semi-persistent manner. Feeds of several hours are needed for the virus to be acquired or transmitted efficiently (Hoyer et al. 1996b; Sim et al. 2000). Gamarra et al. (2010) have reported Bemisia afer sensu lato to be the additional whitefly vector for this virus. The virus also perpetuates through cropping cycles via infected cuttings. The virus is not mechanically sap-transmissible, and aphid vectors failed to transmit this disease (Valverde et al. 2004b). Virion properties and genome The virions are non-enveloped, bipartite filamentous particles about 650–850 nm and 700–900 nm in length, and 10–13 nm in diameter. The genome consists of two linear, positive-sense, single-stranded RNAs which are encapsidated in a single capsid polypeptide of M 43 kd (Koonin and Dolja 1993; Kreuze et al. 2002). The complete nucleotide sequences of SPCSV isolates (serotype East Africa 2) RNA1 (AJ428554 = NC_004123) and RNA2 (AJ428555 = NC_004124) are 9407 nt and 8223 nt, respectively (Alicai et al. 1999; Kreuze et al. 2002; Tairo et al. 2005; Cuellar et al. 2008, 2011b; Kreuze 2011; Qin et al. 2013b; Tugume et al. 2013; Tzanetakis et al. 2013).

Sweet potato collusive virus

(SPCV)

Synonyms Sweet potato caulimo-like virus Taxonomic position Genus: Cavemovirus

Family: Caulimoviridae

Geographical distribution SPCV infection in plants of Ipomoea batatas spreads in the USA, Africa, Puerto Rico, New Zealand, Kenya, Nigeria, Egypt, Papua New Guinea, the Solomon Islands, Australia, Tonga, Ghana, and Madeira, Portugal (Atkey and Brunt 1987; Aritua et al. 2007a; Sossah et al. 2015). Symptoms and host(s) The virus-infected sweet potato plants are usually symptomless but occasionally a few leaves have chlorotic or purple spots. Transmission The virus is graft-transmissible. The virus is not transmissible by mechanical inoculation, not by contact between plants, and not by seed. No vector has been identified. Virion properties and genome The virions are isometric, non-enveloped, and 45–50 nm in diameter with no obvious surface structure. The genome is a monopartite, circular double-stranded DNA of 7723 bp (HQ694978 = NC_015328) (Atkey and Brunt 1987; De Souza and Cuellar 2011; Hohn 2011).

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Sweet potato feathery mottle virus Taxonomic position Genus: Potyvirus

(SPFMV)

Family: Potyviridae

Geographical distribution SPFMV infection in plants of Ipomoea batatas has a very wide distribution and now probably occurs wherever sweet potatoes are grown (Moyer 1986; Lopez and Salazar 1987; Cedano et al. 1989; Kumar et al. 1991; Stobbs et al. 1991; Gibb and Padovan 1993; CABI/EPPO 2003; Usugi et al. 1994; Chavi et al. 1997; Sakai et al. 1997; Souto et al. 2003; Ateka et al. 2004b; Jeeva et al. 2004a; Lozano et al. 2004; Valverde et al. 2004a; Parrella et al. 2006; Jones and Dwyer 2007; Untiveros et al. 2008; Prasanth and Hegde 2008; Tugume et al. 2010b; Hegde et al. 2012; Qiao et al. 2012; Qin et al. 2013a; Kwak et al. 2014; Sossah et al. 2015).

Symptoms and host(s) On sweet potato, leaf symptoms of SPFMVare often inconspicuous or absent. If present, leaf symptoms appear as faint, irregular chlorotic spots occasionally bordered by purplish pigment. The classic irregular chlorotic patterns (feathering) along midribs and faint-to-distinct chlorotic spots, with or without purple margins, occur in some cultivars. Symptom intensity on foliage is influenced by cultivar susceptibility, degree of stress, growth stage, and strain virulence. The “russet crack” strain causes external necrotic lesions or internal cork on certain varieties (Kumar et al. 1991; Karyeija et al. 2001; Jeeva et al. 2004b; Gibson et al. 2014). Campbell et al. (1974) and Cali and Moyer (1981) have reported the etiology of sweet potato russet crack disease which was producing symptoms like necrotic lesions on the fine roots of certain sweet potato cultivars, identified as a strain known as SPFMV-RC.

Transmission The virus is transmitted by aphid vectors Myzus persicae, Aphis gossypii, A. craccivora, and Lipaphis erysimi in a non-persistent manner (Pozzer et al. 1993). The virus is transmissible experimentally by grafting and by mechanical inoculation to its known hosts. The experimental host range of the virus is mainly restricted to species of the Convolvulaceae and Chenopodiaceae; a few strains, however, also infect species of the Solanaceae (Akel et al. 2010). The virus is not transmissible by contact between plants, and the virus is not seed-borne, pollen-borne, or soil-borne. The virus is disseminated in infected tubers and cuttings (Moyer et al. 1980; Valverde et al. 2004b; Parrella et al. 2006).

Virion properties and genome The virions are non-enveloped, flexuous filaments, 810–865 nm long, and 12 nm wide and have helical symmetry with a pitch of ca 3.4 nm (Cali and Moyer 1981). The genome consists of a single molecule of linear, positive-sense ssRNA of 10,820 nt (D86371 = NC_001841) (Moyer and Kennedy 1978; Moyer and Cali 1985; Jeeva et al. 2004a; Sakai et al. 2009; Yamasaki et al. 2010). The genome organization of the isolate Bungo (strain group C) was very different from those of isolates in the russet crack, ordinary (O), and East African groups. 10-O appeared to be a recombinant of isolates S and O, with a recombination site within the P1 gene (Colinet et al. 1994; Sakai et al. 1997; Ryu et al. 1998; Tairo et al. 2005; Yamasaki et al. 2009a; Sharma et al. 2014; Wylie et al. 2017).

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Sweet potato golden vein Korea virus Taxonomic position Genus: Begomovirus

(SPGVKRV)

Family: Geminiviridae

Geographical distribution SPGVKRV infection in plants of Ipomoea batatas was reported from South Korea (Kim et al. unpublished; KT992056). Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2807 nt (KT992056) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Sweet potato latent virus Taxonomic position Genus: Potyvirus

(SPLV)

Family: Potyviridae

Geographical distribution SPLV infection in plants of Ipomoea batatas occurs in Asia, Africa, Japan, Taiwan, China, Korea, and Egypt (Clark and Moyer 1988; Usugi et al. 1991; Makeshkumar et al. 2001; Yun et al. 2002; Qiao et al. 2012; Wassawa 2012; Qin et al. 2013a; Wang et al. 2013a; Kwak et al. 2014). Symptoms and host(s) The virus-infected sweet potato plants exhibit latent or mild chlorosis symptoms. Most Ipomoea batatas cultivars are infected symptomlessly. Transmission The virus is transmitted by the aphid vector, Myzus persicae. The virus is transmissible by mechanical sap-inoculation to many species of the Convolvulaceae family, and to certain Chenopodium spp. and Nicotiana spp. The virus is transmissible by grafting but not transmissible by contact between plants and by seed (Moyer et al. 1989). Virion properties and genome The virions are non-enveloped and flexuous filaments, with a clear modal length of 750–790 nm and 13 nm in wide (Usugi et al. 1991; Wang et al. 2007). The capsid protein has a molecular weight of 36,000. The genome consists of a single molecule of linear, positive-sense ssRNA 10,081 nt (KC443039 = NC_020896) (Wang et al. 2013a). The genome consists of a single large open reading frame encoding a polyprotein of 3247 amino acids (Hammond et al. 1992; Colinet et al. 1997; Revers and Garcia 2015; Wylie et al. 2017).

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Sweet potato leaf curl Canary virus Taxonomic position Genus: Begomovirus

(SPLCCV)

Family: Geminiviridae

Geographical distribution SPLCCV infection in plants of Ipomoea batatas was reported from the Canary Islands, Spain, and China (Lozano et al. 2009; Wassawa 2012; Liu et al. 2017). Symptoms and host(s) The virus-infected sweet potato plants exhibit symptoms of leaf curling, yellowing, and stunting. Transmission Insect tranmsision of SPLCCV has not been demonstrated. However, in common with all other begomoviruses, SPLCCV is likely transmitted by the whitefly vector of begomoviruses, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of SPLCCV has not been investigated. In common with all geminiviruses, the virions of SPLCCV are likely geminate (twinned icosahedra). The genome of SPLCCV consists of a single, circular single-stranded DNA of 2807 nt (FJ529203 = NC_013465; EF456742, EF456745, EU856365) (Briddon 2001; Lozano et al. 2009; Brown et al. 2015; Zerbini et al. 2017). The characterized genome of SPLCCV encodes the six genes typically encoded by monopartite begomoviruses and the DNA-A component of bipartite begomoviruses. The expression and function of the genes have not been investigated for SPLCCV.

Sweet potato leaf curl China virus Taxonomic position Genus: Begomovirus

(SPLCCNV)

Family: Geminiviridae

Geographical distribution SPLCCNV infection in plants of Ipomoea batatas was reported from China (Luan et al. 2007; Bi and Zhang 2012; Wassawa 2012). Symptoms and host(s) The virus-infected sweet potato plants exhibit symptoms of leaf curling, yellowing, foliar mottling, and stunting. Transmission Insect transmission of SPLCCNV has not been demonstrated. However, in common with all other begomoviruses, SPLCCNV is likely transmitted by the whitefly vector of begomoviruses, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of SPLCCNV has not been investigated. In common with all geminiviruses, the virions of SPLCCNV are likely geminate (twinned icosahedra).

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The genome of SPLCCNV consists of a single, circular single-stranded DNA of 2771 nt (DQ512731 = NC_038464; JF736657, JX961671, JX961673, JX961674) (Briddon 2001; Luan et al. 2007; Brown et al. 2015; Zerbini et al. 2017). The characterized genome of SPLCCNV encodes the six genes typically encoded by monopartite begomoviruses and the DNA-A component of bipartite begomoviruses. The expression and function of the genes have not been investigated for SPLCCNV.

Sweet potato leaf curl Georgia virus

(SPLCGV)

Synonyms Ipomoea leaf curl virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution SPLCGV infection in plants of Ipomoea batatas was reported from Argentina, Georgia (USA), China, India, and South Africa (Lotrakul et al. 2003; Prasanth and Hegde 2008; Liu et al. 2013; Qin et al. 2013c; Kashif et al. 2012; Wassawa 2012; Esterhuizen et al. 2012; Hegde et al. 2012; Martino et al. 2017). Symptoms and host(s) The virus-infected sweet potato plants exhibit symptoms of upward leaf curling, leaf rolling, chlorosis, and stunting. Transmission SPLCGV is transmitted by the whitefly vector of begomoviruses, Bemisia tabaci, in a circulative, nonpropagative manner (Lotrakul et al. 2003). Virion properties and genome The structure of the virions of SPLCGV has not been investigated. In common with all geminiviruses, the virions of SPLCGV are likely geminate (twinned icosahedra). The genome of SPLCGV consists of a single, circular single-stranded DNA of 2773 nt (AF326775 = NC_004640; JX448368) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). The characterized genome of SPLCGV encodes the six genes typically encoded by monopartite begomoviruses and the DNA-A component of bipartite begomoviruses. The expression and function of the genes have not been investigated for SPLCGV.

Sweet potato leaf curl Guangxi virus Synonyms Sweet potato leaf curl china virus 2 Taxonomic position Genus: Begomovirus

Family: Geminiviridae

(SPLCGV)

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Geographical distribution SPLCGV infection in plants of Ipomoea batatas was reported from China (Liu et al. 2017). Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2831 nt (KJ476510 = NC_024693) (Briddon 2001; Brown et al. 2015; Liu et al. 2017; Zerbini et al. 2017).

Sweet potato leaf curl Henan virus Taxonomic position Genus: Begomovirus

(SPLCHnV)

Family: Geminiviridae

Geographical distribution SPLCHnV infection in plants of Ipomoea batatas was detected in Henan Province, China (Liu et al. 2014). Symptoms and host(s) The virus-infected sweet potato plants exhibit severe leaf curl symptoms. Transmission The transmission of SPLCHnV has not been investigated. It is likely that, in common with other begomoviruses, SPLCHnV will be transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Similarly it is likely that the virus will prove not to be either mechanically or seedtransmitted. Virion properties and genome The structure of the virions of SPLCHnV has not been investigated. In common with all geminiviruses, the virions of SPLCHnV are likely geminate (twinned icosahedra). With only a single isolate of SPLCHnV having been characterized, the precise nature of the genome (whether monopartite, monopartite betasatellite-associated, or bipartite) is unclear. However, the majority of sweet-potato-infecting begomoviruses are monopartite. The characterized genome/genomic component of SPLCHnV is 2766 nt (KC907406 = NC_021719) (Briddon 2001; Liu et al. 2014; Brown et al. 2015; Zerbini et al. 2017). The characterized genome of SPLCHnV encodes the six genes typically encoded by monopartite begomoviruses. The expression and function of the genes have not been investigated.

Sweet potato leaf curl Sao Paulo virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

(SPLCSPV)

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Geographical distribution SPLCSPV infection in plants of Ipomoea batatas was reported from Uganda, Brazil, and South Africa (Albuquerque et al. 2011, 2012; Esterhuizen et al. 2012). Symptoms and host(s) The virus-infected sweet potato plants exhibit symptoms of leaf curling, yellowing, and stunting. Transmission Insect transmission of SPLCSPV has not been demonstrated. However, in common with all other begomoviruses, SPLCSPV is likely transmitted by the whitefly vector of begomoviruses, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of SPLCSPV has not been investigated. In common with all geminiviruses, the virions of SPLCSPV are likely geminate (twinned icosahedra). The genome of SPLCSPV consists of a single, circular single-stranded DNA of 2782 nt (HQ393477 = NC_025681; JQ621844) (Briddon 2001; Albuquerque et al. 2011, 2012; Brown et al. 2015; Zerbini et al. 2017). The characterized genome of SPLCSPV encodes the six genes typically encoded by monopartite begomoviruses and the DNA-A component of bipartite begomoviruses. The expression and function of the genes have not been investigated for SPLCSPV.

Sweet potato leaf curl Shandong virus Taxonomic position Genus: Begomovirus

(SPLCSdV)

Family: Geminiviridae

Geographical distribution SPLCSdV infection in plants of Ipomoea batatas was reported from China (Wang and Zhang, unpublished-KU323597) Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22 × 38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2826 nt (KU323597) (Brown et al. 2015; Zerbini et al. 2017).

Sweet potato leaf curl Sichuan virus 1 Taxonomic position Genus: Begomovirus

Family: Geminiviridae

(SPLCSiV-1)

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Geographical distribution SPLCSiV-1 infection in plants of Ipomoea batatas was reported from Henan Sichuan, China (Liu et al. 2014). The virus was previously known as Sweet potato leaf curl China Sichuan virus (Liu et al. 2013; Brown et al. 2015). Symptoms and host(s) The virus-infected sweet potato plants exhibit leaf curling and yellowing symptoms. Transmission The transmission of SPLCSiV-1 has not been investigated. It is likely that, in common with other begomoviruses, SPLCSiV-1 will be transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Similarly it is likely that the virus will prove not to be either mechanically or seedtransmitted. Virion properties and genome The structure of the virions of SPLCSiV-1 has not been investigated. In common with all geminiviruses, the virions of SPLCSiV-1 are likely geminate (twinned icosahedra). With only a single isolate of SPLCSiV-1 having been characterized, the precise nature of the genome (whether monopartite, monopartite betasatellite-associated, or bipartite) is unclear. However, the majority of sweet-potato-infecting begomoviruses are monopartite. The characterized genome/ genomic component of SPLCSiV-1 is 2764 nt (KC488316 = NC_038465) (Briddon 2001; Liu et al. 2013; Brown et al. 2015; Zerbini et al. 2017). The characterized genome of SPLCSiV-1encodes the six genes typically encoded by monopartite begomoviruses native to the Old World. The expression and function of the genes have not been investigated.

Sweet potato leaf curl Sichuan virus 2 Taxonomic position Genus: Begomovirus

(SPLCSiV-2)

Family: Geminiviridae

Geographical distribution SPLCSiV-2 infection in plants of Ipomoea batatas was reported from China (Brown et al. 2015; Liu et al. 2017). Symptoms and host(s) The virus-infected sweet potato plants exhibit leaf curling and yellowing symptoms. Transmission The transmission of SPLCSiV-2 has not been investigated. It is likely that, in common with other begomoviruses, SPLCSiV-2 will be transmitted by the whitefly Bemisia tabaci in a circulative, nonpropagative manner. Similarly it is likely that the virus will prove not to be either mechanically or seedtransmitted. Virion properties and genome The structure of the virions of SPLCSiV-2 has not been investigated. In common with all geminiviruses, the virions of SPLCSiV-2 are likely geminate (twinned icosahedra).

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With only a single isolate of SPLCSiV-2 having been characterized, the precise nature of the genome (whether monopartite, monopartite betasatellite-associated, or bipartite) is unclear. However, the majority of sweet-potato-infecting begomoviruses are monopartite. The characterized genome/ genomic component of SPLCSiV-2 is 2786 nt (KF156759 = NC_022586) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). The characterized genome of SPLCSiV-2 encodes the six genes typically encoded by monopartite begomoviruses native to the Old World. The expression and function of the genes have not been investigated.

Sweet potato leaf curl South Carolina virus Taxonomic position Genus: Begomovirus

(SPLCSCV)

Family: Geminiviridae

Geographical distribution SPLCSCV infection in plants of Ipomoea batatas was reported from South Carolina, USA (Zhang and Ling 2011). Symptoms and host(s) The virus-infected sweet potato plants exhibit symptoms of leaf curling, yellowing, and stunting. Transmission Insect transmission of SPLCSCV has not been demonstrated. However, in common with all other begomoviruses, SPLCSCV is likely transmitted by the whitefly vector of begomoviruses, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of SPLCSCV has not been investigated. In common with all geminiviruses, the virions of SPLCSCV are likely geminate (twinned icosahedra). The genome of SPLCSCV consists of a single, circular single-stranded DNA of 2782 nt (HQ333144 = NC_015317) (Briddon 2001; Zhang and Ling 2011; Brown et al. 2015; Zerbini et al. 2017). The characterized genome of SPLCSCVencodes the six genes typically encoded by monopartite begomoviruses and the DNA-A component of bipartite begomoviruses. The expression and function of the genes have not been investigated for SPLCSCV.

Sweet potato leaf curl virus

(SPLCV)

Synonyms Ipomoea yellow vein virus (IYVV) Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution SPLCV infection in plants of Ipomoea batatas has an almost global distribution, having been reported from almost all areas where sweet potato (Ipomoea batatas) is grown including India, Argentina, Israel, Japan, Brazil, Puerto Rico, South Korea, China, Spain, Kenya, Peru, Sicily, Italy, Sudan, and the USA

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(Osaki and Inouye 1991; Lotrakul et al. 1998, 2002, 2003; Lotrakul 2000; Fuentes and Salazar 2003; Li et al. 2004; Briddon et al. 2006; Luan et al. 2006; Miano et al. 2006; Makeshkumar et al. 2007; Lozano et al. 2009; Park et al. 2011; Hegde et al. 2012; Pardina et al. 2012; Qiao et al. 2012; Kwak et al. 2014; Inoue-Nagata et al. 2016; Liu et al. 2017; Mohammed et al. 2017). Various isolates of this virus have previously been known under the names Sweet potato leaf curl Spain virus, Sweet potato leaf curl Lanzarote virus, Sweet potato golden vein-associated virus, Sweet potato leaf curl Bengal virus, and Sweet potato leaf curl Shanghai virus (Kil et al. 2014). Symptoms and host(s) The virus-infected sweet potato plants produce upward curling of leaves and vein swelling on young plants but later few or no symptoms. The natural host range was limited to plants in the genus Ipomoea within the family Convolvulaceae (Ling et al. 2011). The virus has been isolated from sweet potato (Ipomoea batatas), I. indica, I. purpurea, and I. setosa. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. A minimum of 15 min was required for a viruliferous whitefly population to transmit the virus to I. setosa. The efficiency of transmission increased over time; however, only 60% of the assay plants were infected after exposure to viruliferous whiteflies for 48 h. Male and female adult whiteflies vectored SPLCV with similar efficiency (Chung et al. 1985; Valverde et al. 2004c; Simmons et al. 2009). The virus is not transmissible by mechanical inoculation, by grafting, and by contact between plants. SPLCV has been detected in >70% of the seeds from the infected sweet potato plants and the seed-transmission up to 15% was obtained, which demonstrated the first evidence of seed-transmission in geminivirus (Kim et al. 2015). Virion properties and genome The structure of the virions of SPLCV has not been investigated. In common with all geminiviruses, the virions of SPLCV are likely geminate (twinned icosahedra). The genome of SPLCV consists of a single, circular single-stranded DNA of 2828 nt (AF104036 = NC_004650; AJ132548) (Lotrakul et al. 1998; Lotrakul and Valverde 1999; Briddon 2001; Luan et al. 2007; Lozano et al. 2009; Trenado et al. 2011; Pardina et al. 2012; Brown et al. 2015; Zerbini et al. 2017). The characterized genome of SPLCV encodes the six genes typically encoded by monopartite begomoviruses and the DNA-A component of bipartite begomoviruses. The expression and function of the genes have not been investigated for SPLCV. Three deltasatellites are associated with SPLCV, viz., sweet potato leaf curl deltasatellite 1 (662 nt; FJ914390), sweet potato leaf curl deltasatellite 2 (733 nt; KF716173), and sweet potato leaf curl deltasatellite 3 (731 nt; KT099179) (Zhou 2013; Hassan et al. 2016; Lozano et al. 2016; Rosario et al. 2016).

Sweet potato leaf speckling virus Taxonomic position Genus: Unassigned

(SPLSV)

Family: Luteoviridae

Geographical distribution SPLSV infection in plants of Ipomoea batatas was reported from Peru and Cuba (Fuentes et al. 1996).

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Symptoms and host(s) The virus-infected sweet potato plants show leaf curling and clear whitish flecks or specks on the leaves (Fuentes et al. 1996). Transmission The virus is transmitted by an aphid vector Macrosiphum euphorbiae in a persistent manner, whereas Myzus persicae and Aphis gossypii failed to transmit the virus. It infects 45 species of 14 plant families (Moyer and Salazar 1989), including a number of Ipomoea species. The virus is not mechanically sap-transmissible. Virion properties and genome The virions are isometric c. 30 nm in diameter. SPLSV particles did not react with antibodies to PLRV in ISEM or ELISA tests. Only the coat protein gene sequence of SPLSV is available (DQ655700), which shares 70% identity with that of PLRV (Fuentes et al. 1996).

Sweet potato mild mottle virus Taxonomic position Genus: Ipomovirus

(SPMMV)

Family: Potyviridae

Geographical distribution SPMMV infection in plants of Ipomoea batatas was reported from Africa (Egypt, Burundi, Tanzania, Rwanda, Ghana, Kenya, and Uganda), Indonesia, China, Philippines, Papua New Guinea, New Zealand, and India (Hollings et al. 1976; Makeshkumar et al. 2001; Mukasa et al. 2003a; Ateka et al. 2004a; Tairo et al. 2004; Njeru et al. 2008; Nyaboga et al. 2008; Tugume et al. 2010a; Kathurima et al. 2011; Hegde et al. 2012; Sossah et al. 2015). Symptoms and host(s) The virus-infected sweet potato plants exhibit symptoms of vein-clearing, leaf mottling, and stunting. Some sweet potato cultivars were symptomless. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a non-persistent manner (Hollings et al. 1976; Jones 2003; Dombrovsky et al. 2014; Gilbertson et al. 2015). The virus is transmissible by mechanical sap-inoculation, and has wide host range of herbaceous plant species which includes 45 species in 14 families. In certain species of tobacco and Chenopodium, locals lesions were produced. The virus is transmissible by grafting. The virus is not transmissible by contact between plants and not by seed. Virion properties and genome The virions are non-enveloped, flexuous filaments, 800–950 nm long, and 12–15 nm in diameter. The genome consists of a single molecule of linear, positive-sense ssRNA of 10,818 nt (Z73124 = NC_003797) and contains a polyadenylated tract at the 30 terminus. The coat protein has a molecular weight of ~35 kDa (Hollings et al. 1976; Colinet et al. 1998a; Mukasa et al. 2003b; Tairo et al. 2005; Lopez-Moya et al. 2009; Colinet 2011; Wylie et al. 2017).

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Sweet potato mild speckling virus Taxonomic position Genus: Potyvirus

(SPMSV)

Family: Potyviridae

Geographical distribution SPMSV infection in plants of Ipomoea batatas occurs in Peru, Argentina, China, South Africa, Nigeria, New Zealand, the Philippines, Egypt, Ghana, and Indonesia (Alvarez et al. 1997; Nome et al. 2006; Untiveros et al. 2008; Sossah et al. 2015). Symptoms and host(s) The virus-infected sweet potato plants exhibit chlorosis, dwarfing, vein-clearing, and leaf distortion symptoms. The host range of the virus is restricted to species of the families Convolvulaceae, Chenopodiaceae, and Solanaceae. Transmission The virus is transmitted by an aphid vector Myzus persicae, in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, 800 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA. Only partial sequence of SPMSV containing 1103 nt representing the coat protein cistron and 30 noncoding region (U61228) is available (Alvarez et al. 1997; Nome et al. 2006; Revers and Garcia 2015; Wylie et al. 2017).

Sweet potato mosaic virus Taxonomic position Genus: Begomovirus

(SPMV)

Family: Geminiviridae

Geographical distribution SPMV infection in plants of Ipomoea batatas was reported from Brazil (Paprotka et al. 2010; Wassawa 2012). Symptoms and host(s) The virus-infected sweet potato plants exhibit mosaic symptoms. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminated) incomplete icosahedra, T = 1, 22  38 nm, with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2803 nt (FJ96983 = NC_038466) (Briddon 2001; Paprotka et al. 2010; Brown et al. 2015; Zerbini et al. 2017).

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Sweet potato pakakuy virus

(SPPV)

Synonyms Sweet potato badnavirus A (SPBV-A); Sweet potato badnavirus B (SPBV-B) Taxonomic position Genus: Badnavirus

Family: Caulimoviridae

Geographical distribution SPPV infection in plants of Ipomoea batatas was reported from Peru, Tanzania, China, and South Africa (Kreuze et al. 2009; Mbanzibwa et al. 2011; Kashif et al. 2012; Qin et al. 2016; Nhlapo et al. 2018), and have since been reported in germplasm and field material from many countries (Kreuze et al. 2017). Symptoms and host(s) The virus-infected sweet potato plants exhibit vein-clearing and mosaic symptoms. Transmission The virus is not mechanically sap-transmissible and not transmissible by contact between plants. The virus is transmitted efficiently through seed (Kreuze et al. 2017). The use of virus-infected vegetative planting material is the main mode of virus spread. Virion properties and genome The virions are bacilliform in shape with a diameter of 30 nm and modal particle length of 130 nm. The genome is a monopartite, circular double-stranded DNA of about 8 kbp with a single-strand discontinuity at one site in each strand; strains are variable, and two full genomes are available (FJ560943, 8082 bp, ‘SPPV’; FJ560944, 7961 bp ‘SPVB-B’) (Olszewski and Lockhart 2011; Bhat et al. 2016; Kreuze et al. 2017).

Sweet potato symptomless virus 1 Taxonomic position Genus: Mastrevirus

(SpSV-1)

Family: Geminiviridae

Geographical distribution SpSV-1 infection in plants of Ipomoea batatas was reported from Kenya, China, Peru, Tanzania, Taiwan, Uruguay, the USA and several Central American countries (Mbanzibwa et al. 2011; Clark et al. 2012; Kashif et al. 2012; Wang et al. 2015; Cao et al. 2017). Symptoms and host(s) The virus-infected plants exhibit mild mottling or mosaic symptoms. Transmission The virus is transmitted by the leafhopper vectors in a persistent, circulative and non-propagative manner. The virus is not transmitted by mechanical inoculation.

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Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is a monopartite circular single-stranded DNA of 2602 nt (KY565231). There are coding regions in both the virion (positive) and complementary (negative) sense strands (Cao et al. 2017; Zerbini et al. 2017).

Sweet potato vein clearing virus Taxonomic position Genus: Solendovirus

(SPVCV)

Family: Caulimoviridae

Geographical distribution SPVCV detected in plants of Ipomoea batatas was from East Africa, Central America and Fiji, but not in South America (Cuellar et al. 2011a; Wassawa 2012; Wu et al. 2018). Symptoms and host(s) SPVCV causes vein-clearing symptoms in the indicator plant, Ipomoea setosa. In sweet potato it causes symptomless infection. Transmission The virus is not sap-transmissible. The virus is readily transmissible by grafting to sweet potato and I. setosa plants with 100% efficiency. No vector has been identified. Virion properties and genome The virions are isometric, non-enveloped, and 50 nm in diameter with no obvious surface structure. The genome is a monopartite, circular double-stranded DNA of 8837 bp (HQ694979 = NC_015228) (Cuellar et al. 2011a).

Sweet potato virus 2

(SPV2)

Synonyms Sweet potato vein mosaic virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution SPV2 infection in plants of Ipomoea batatas was reported from Taiwan, North Carolina (USA), China, Spain, South Africa, Portugal, Australia, Barbados, Peru, New Zealand, Korea, and Zambia (Nome 1973; Souto et al. 2003; Ateka et al. 2004a, 2007; Tairo et al. 2006; Trenado et al. 2007; Untiveros et al. 2008; Perez-Egusquiza et al. 2009; Almeyda et al. 2013; Qin et al. 2013a; Kwak et al. 2014). Symptoms and host(s) The virus-infected sweet potato plants exhibit mild symptoms of leaf mottle, vein yellowing, and ringspots (Almeyda et al. 2013).

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Transmission The virus is transmitted by two biotypes of Myzus persicae in a non-persistent manner. The virus is mechanically transmissible to several species of the genera Chenopodium, Datura, Nicotiana, and Ipomoea (Ateka et al. 2004a). Virion properties and genome The virions are non-enveloped, flexuous filaments, 850 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 10,731 nt (JN613807 = NC_017970) (Ateka et al. 2004a, 2007; Li et al. 2012; Revers and Garcia 2015; Wylie et al. 2017).

Sweet potato virus C Taxonomic position Genus: Potyvirus

(SPVC)

Family: Potyviridae

Geographical distribution SPVC was previously reported as SPFMV-C strain or SPFMV-Bungo isolate in Japan (Sakai et al. 2009). SPVC was also found affecting plants of Ipomoea batatas in Israel, China, Korea, Honduras, and Guatemala (Clark et al. 2012; Qin et al. 2013a; Prakash et al. 2013; Kashif et al. 2012; Kwak et al. 2014). Symptoms and host(s) The virus-infected sweet potato plants were of poor appearance, grew feebly, and showed strong leaf malformation, mosaic, purpling, chlorosis, cupping, vein yellowing, and occasional feathery mottle symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is transmissible through mechanical sap-inoculation, and grafting. Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide. The genome is a single molecule of linear positive-sense single-stranded RNA of 10820 nt (GU207957 = NC_014742) (Yamasaki et al. 2010; Revers and Garcia 2015; Wylie et al. 2017).

Sweet potato virus G Taxonomic position Genus: Potyvirus

(SPVG)

Family: Potyviridae

Geographical distribution SPVG infection in plants of Ipomoea batatas was first reported from China (Colinet et al. 1998b) and subsequently from New Zealand, Hawaii, French Polynesia, Easter Island, Africa (Egypt, Ethiopia, Ghana, Nigeria, and Zimbabwe), Eastern Asia (China, Japan, Java,Korea, and Taiwan), the southern USA, South America (Peru), Europe (Spain), and Barbados (IsHak et al. 2003; Souto et al. 2003; Alemu 2004; Trenado et al. 2007; Rannali et al. 2008; Untiveros et al. 2008; Yamasaki et al. 2009b;

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Qiao et al. 2012; Almeyda et al. 2013; Qin et al. 2013a; Wang et al. 2013b; Kwak et al. 2014; Sossah et al. 2015). Symptoms and host(s) The virus-infected sweet potato plants showed virus-like symptoms including mottling, vein-clearing, chlorotic mosaic, and chlorotic spots (Yamasaki et al. 2009b). Transmission The virus is transmitted by aphid vectors Myzus persicae and Aphis gossypii in a non-persistent manner. The virus is also mechanically sap-transmissible (Gu et al. 2006). Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long, and 11–13 nm wide, with helical symmetry and a pitch of about 3.4 nm. The genome consists of a single molecule of linear, positivesense single-stranded RNA of 10,798 nt (JQ824374 = NC_018093, JN613805) (Gu et al. 2006; Li et al. 2012; Rodriguez Pardina et al. 2012; Revers and Garcia 2015; Wylie et al. 2017).

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Virus Res 49:91–100 Colinet D, Kummert J, Lepoivre P (1998a) The nucleotide sequence and genome organization of the whitefly transmitted sweet potato mild mottle virus: a close relationship with members of the family Potyviridae. Virus Res 53:187–196 Colinet D, Nguyen M, Kummert J, Lepoivre P, Xia FZ (1998b) Differentiation among potyviruses infecting sweet potato based on genus- and virus-specific reverse transcription polymerase chain reaction. Plant Dis 82:223–229 Cuellar WJ, Tairo F, Kreuze JF, Valkonen JP (2008) Analysis of gene content in sweet potato chlorotic stunt virus RNA1 reveals the presence of the p22 RNA silencing suppressor in only a few isolates: implications for viral evolution and synergism. J Gen Virol 89(2):573–582 Cuellar WJ, De Souza J, Barrantes I, Fuentes S, Kreuze JF (2011a) Distinct cavemoviruses interact synergistically with sweet potato chlorotic stunt virus (genus Crinivirus) in cultivated sweet potato. 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Dombrovsky A, Reingold V, Antignus Y (2014) Ipomovirus – an atypical genus in the family Potyviridae transmitted by whiteflies. Pest Manag Sci 70(10):1553–1567 Esterhuizen L, van Heerden S, Rey M, van Heerden H (2012) Genetic identification of two sweet-potato-infecting begomoviruses in South Africa. Arch Virol 157(11):2241–2245 Fuentes S (1994) Preliminary identification of a sweet potato virus (C-6). Fitopatologia 29:38 Fuentes S, Salazar LF (1989) Identification of sweet potato (Ipomoea batatas (L.) Lam) viruses. Fitopatología 24:43 Fuentes S, Salazar LF (1992) Identification of a new sweet potato virus. Fitopatologia 27:50 Fuentes S, Salazar LF (2003) First report of Sweet potato leaf curl virus in Peru. Plant Dis 87:98 Fuentes S, Mayo MA, Jolly CA, Nakano M, Querci M, Salazar LF (1996) A novel luteovirus from sweet potato, sweet potato leaf speckling virus. 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Ann Appl Biol 82:511–528 Hoyer U, Maiss E, Jelkmann W, Lesemann D-E, Vetten HJ (1996a) Identification of the coat protein gene of a sweet potato sunken vein closterovirus isolate from Kenya and evidence for serological relationship among geographically diverse closterovirus isolates from sweet potato. Phytopathology 86:744–750 Hoyer U, Jelkmann W, Maiss E, Vetten HJ (1996b) Sweet potato sunken vein virus (SPSVV): another bipartite closterovirus transmitted by Bemisia tabaci. In Abstracts of Presentations on Plant Protection Issues, Xth International Congress of Virology, Jerusalem, Israel. Phytoparasitica 26, issue 1. http://www.phytoparasitica.org/phyto/org Inoue-Nagata AK, Lima MF, Gilbertson RL (2016) A review of geminivirus (Begomovirus) diseases in vegetables and other crops in Brazil: current status and approaches for management. Hortic Bras 34:8–18 IsHak JA, Kreuze JF, Johansson A, Mukasa SB, Tairo F, Abo El-Abbas FM, Valkonen JPT (2003) Some molecular characteristics of three viruses from SPVD affected sweet potato plants in Egypt. Arch Virol 148:2449–2460 Jeeva ML, Balakrishnan S, Edison S, Umamaheswaran K, Makeshkumar T (2004a) Characterization, purification and serology of Sweet potato feathery mottle virus in India. J Root Crops 30(1):24–30 Jeeva ML, Balakrishnan S, Edison S (2004b) Symptomatology and host range of sweet potato feathery mottle disease. J Mycol Pl Pathol 34:180–184 Jones DR (2003) Plant viruses transmitted by whiteflies. Eur J Plant Pathol 109:195–219 Jones RAC, Dwyer GI (2007) Detection of Sweet potato chlorotic fleck virus and Sweet potato feathery mottle virus – strain O in Australia. Aust Plant Pathol 36:591–594 Karyeija RF, Kreuze JF, Gibson RW, Valkonen JPT (2001) Variability of Sweet potato feathery mottle virus in Africa. 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Kathurima TM, Bett BB, Miano DW, Kim DJ (2011) Diagnostics of viruses infecting local farmer preferred sweet potato cultivars in Kenya. Afr J Agric Res 6:3718–3724 Kil EJ, Kim J, Byun HS, Kim S, Kwak HR, Kim MK, Choi HS, Chung MN, Lee S (2014) First report of sweet potato golden vein associated virus infecting sweet potato in Korea. Plant Dis 98(8):116 Kim J, Kil E-J, Kim S, Seo H, Byun H-S, Park J, Chung M-N, Kwak H-R, Kim M-K, Kim C-S, Yang J-W, Lee K-Y, Choi H-S, Lee S (2015) Seed transmission of sweet potato leaf curl virus in sweet potato (Ipomoea batatas). Plant Pathol 64:1284–1291 Koonin EV, Dolja VV (1993) Evolution and taxonomy of positive strand RNA viruses: implications of comparative analysis of amino acid sequences. Crit Rev Biochem Mol Biol 28:375–430 Kreuze JF (2011) Crinivirus. Closteroviridae. The Springer Index of Viruses. 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Curr Sci 60:321–325 Kwak H-R, Kim M-K, Shin J-C, Lee Y-J, Seo J-K, Lee H-U, Jung M-N, Kim S-H, Choi H-S (2014) The current incidence of viral disease in Korean sweet potatoes and development of multiplex RT-PCR assays for simultaneous detection of eight sweet potato viruses. Plant Pathol J 30:416–424 Li R, Salih S, Hurtt S (2004) Detection of geminiviruses in sweet potato by polymerase chain reaction. Plant Dis 88:1347–1351 Li F, Xu D, Abad J, Li R (2012) Phylogenetic relationships of closely related potyviruses infecting sweet potato determined by genomic characterization of Sweet potato virus G and Sweet potato virus 2. Virus Genes 45(1):118–125 Ling K, Harrison HF Jr, Simmons AM, Zhang S, Jackson DM (2011) Experimental host range and natural reservoir of Sweet potato leaf curl virus in the United States. Crop Prot 30:1055–1062 Liu Q, Zhang Z, Qiao Q, Qin Y, Zhang D, Tian Y, Wang S, Wang Y (2013) Complete genome sequence of a novel monopartite begomovirus infecting sweet potato in China. Virus Genes 47(3):591–594 Liu Q, Zhang Z, Li J, Qiao Q, Qin Y, Zhang D, Tian Y, Wang S, Wang Y (2014) Complete genome sequence of a novel monopartite begomovirus infecting sweet potato in China. Arch Virol 159(6):1537–1540 Liu Q, Wang Y, Zhang Z, Lv H, Qiao Q, Qin Y, Zhang D, Tian Y, Wang S, Li J (2017) Diversity of sweepoviruses infecting sweet potato in China. Plant Dis 101:2098–2103 Lopez D, Salazar LF (1987) Studies on Sweet potato feathery mottle virus (SPFMV) in Peru. Fitopatologı’a 22:40 Lopez-Moya JJ, Valli A, Garcia JA (2009) Potyviridae. In: Encyclopaedia of Life Sciences (ELS). Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000755.pub2 Lotrakul P (2000) Biological and molecular properties of Sweet potato leaf curl virus. Ph.D. Thesis. Louisiana State University. Baton Rouge 93 p Lotrakul P, Valverde RA (1999) Cloning of a DNA-A-like genomic component of sweet potato leaf curl virus: nucleotide sequence and phylogenetic relationships. Mol Plant Pathol On-Line 28: Available: http://www.bspp.org.uk/mppol/ 1999/0422lotrakul Lotrakul P, Valverde RA, Clark CA, Sim J, De La Torre R (1998) Detection of a geminivirus infecting sweet potato in the United States. Plant Dis 85:1253–1257 Lotrakul P, Valverde RA, Clark CA, Hurtt S, Hoy MW (2002) Sweet potato leaf curl virus and related geminiviruses in sweet potato. Acta Hortic 583:135–141 Lotrakul P, Valverde RA, Clark CA, Fauquet CM (2003) Properties of a Begomovirus isolated from sweet potato [Ipomoea batatas (L.) Lam.] infected with sweet potato leaf curl virus. Rev Mex Fitopatología 21:128–136 Lozano G, Navas-Castillo J, Ramos A, Valdes F (2004) First report of Sweet potato chlorotic stunt virus and Sweet potato feathery mottle virus infecting sweet potato in Spain. Plant Dis 88:428 Lozano G, Trenado HP, Valverde RA, Navas-Castillo J (2009) Novel begomovirus species of recombinant nature in sweet potato (Ipomoea batatas) and Ipomoea indica: taxonomic and phylogenetic implications. J Gen Virol 90: 2550–2562 Lozano G, Trenado HP, Fiallo-Olivé E, Chirinos D, Geraud-Pouey F, Briddon RW, Navas-Castillo J (2016) Characterization of non-coding DNA satellites associated with sweepoviruses (genus Begomovirus, Geminiviridae) – definition of a distinct class of begomovirus-associated satellites. Front Microbiol 7:162 Luan YS, Zhang J, An L (2006) First report of Sweet potato leaf curl virus in China. Plant Dis 90:1111

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Luan YS, Zhang J, Liu DM, Li WL (2007) Molecular characterization of sweet potato leaf curl virus isolate from China (SPLCV-CN) and its phylogenetic relationship with other members of the Geminiviridae. Virus Genes 35(2):379–385 Makeshkumar T, Sriram S, Nayar RR, Edison S, Thankappan M (2001) Symptomatology and characterization of sweet potato virus diseases. J Root Crops 27:296–302 Makeshkumar T, Prakash Krishnan BS, Hegde V, Jeeva ML, Edison S (2007) Sweet potato leaf curl disease – a new emerging virus problem of sweet potato in India. In: 10th International Sympoium, October 2007, ICRISAT, Hyderabad India Martin WJ (1962) Susceptibility of certain Convolvulaceae to internal cork tobacco ring spot and cucumber mosaic viruses. Phytopathology 52:607–611 Martino JM, Fontenele RS, Ferreira FA, Ribeiro G, Di Feo LDV (2017) First report of sweet potato leaf curl Georgia virus in sweet potato in Argentina. Plant Dis 101:513 Mbanzibwa DR, Tairo F, Gwandu C, Kullaya A, Valkonen JPT (2011) First report of Sweet potato symptomless virus 1 and sweet potato virus A in Sweet potatoes in Tanzania. Plant Dis 95:224 Miano DW, LaBonte DR, Clark CA, Valverde RA, Hoy MW, Hurtt S, Li R (2006) First report of a begomovirus infecting sweet potato in Kenya. Plant Dis 90:832 Milgram M, Cohen J, Loebenstein G (1996) Effects of sweet potato feathery mottle virus and sweet potato sunken vein virus on sweet potato yields and rate of reinfection on virus-free planting material in Israel. Phytoparasitica 24:189–193 Mohammed HS, El Siddig MA, El Hussein AA, Ibrahim FA, Navas-Castillo J, Fiallo-Olive E (2017) First report of sweet potato leaf curl virus infecting sweet potato in Sudan. Plant Dis 101:849 Moyer JF (1986) Variability among strains of Sweet potato feathery mottle virus. Phytopathology 76:1126 Moyer JW, Cali BB (1985) Properties of sweet potato feathery mottle virus RNA and capsid protein. J Gen Virol 65:1185–1189 Moyer JW, Kennedy GG (1978) Purification and properties of Sweet potato feathery mottle virus. Phytopathology 76:1126 Moyer JW, Salazar LF (1989) Viruses and virus-like diseases of sweet potato. Plant Dis 73:451–455 Moyer JW, Cali BB, Kennedy GG, Abou-Ghadir MF (1980) Identification of two Sweet potato feathery mottle virus strains in North Carolina. Plant Dis 64(8):762–764 Moyer JW, Jackson GVH, Frison EA (eds) (1989) FAO/IBPGR technical guidelines for the safe movement of sweet potato germplasm. Food and Agriculture Organization of the United Nations, Rome/International Board for Plant Genetic Resources, Rome Mukasa SB, Rubaihayo PR, Valkonen JPT (2003a) Incidence of viruses and virus like diseases of sweet potato in Uganda. Plant Dis 87:329–335 Mukasa SB, Rubaihayo PR, Valkonen JP (2003b) Sequence variability within the 30 -proximal part of the Sweet potato mild mottle virus genome. Arch Virol 148:487–496 Ngeve JM (1990) Yield stability and yield depression in sweet potato cultivars susceptible to sweet potato virus disease. J Hortic Sci 65:225–230 Nhlapo TF, Mulabisana JM, Odeny DA, Rey MEC, Rees DJG (2018) First report of Sweet potato badnavirus A and Sweet potato badnavirus B in South Africa. Plant Dis 102(9):1865 Njeru RW, Bagabe MC, Nkezabahizi D, Kayiranga D, Kajuga J, Butare L, Ndirigue J (2008) Viruses infecting sweet potato in Rwanda: occurrence and distribution. Ann Appl Biol 153:215–221 Nome SF (1973) Sweet potato vein mosaic in Argentina. Phytopathol Z 77:44–54 Nome CF, Laguna IG, Nome SF (2006) Cytological alterations produced by Sweet potato mild speckling virus. J Phytopathol 154:504–507 Nyaboga EN, Ateka EM, Bulimo WD (2008) Serological detection of virus diseases of sweet potato in Kenya. J Appl Biosc 7:222–229 Olszewski NE, Lockhart B (2011) Badnavirus. Caulimoviridae. The Springer Index of Viruses. Springer, New York, p 263–269. https://doi.org/10.1007/978-0-387-95919-1_40 Opiyo SA, Ateka EM, Owuor PO, Manguro LOA, Karuri HW (2010) Survey of sweet potato viruses in Western Kenya and detection of Cucumber mosaic virus. J Plant Pathol 92:797–801 Osaki T, Inouye T (1991) Transmission characteristics and cytopathology of a whitefly-transmitted virus isolated from sweet potato leaf curl disease. Bull Osaka Pref Univ Ser B 43:17–19 Paprotka T, Boiteux LS, Fonseca ME, Resende RO, Jeske H, Faria JC, Ribeiro SG (2010) Genomic diversity of sweet potato geminiviruses in a Brazilian germplasm bank. Virus Res 149(2):224–233 Pardina PR, Luque A, Nome C, Lopez Colomba E, Fuentes Delgado S, Di Feo L (2012) First report of Sweet potato leaf curl virus infecting sweet potato in Argentina. Aust Plant Dis Notes 7:157–160 Park J, Kim S, Choi E, Kwak H-R, Kim M-K, Lee K-Y, Choi H-S, Lee S (2011) Molecular characterization of sweet potato leaf curl virus (SPLCV) isolates from Korea: phylogenetic relationship and recombination analysis. Acta Virol 55(4):327–335

Ipomoea batatas (Sweet potato)

1269

Parrella G, De Stradis A, Giorgini M (2006) Sweet potato feathery mottle virus is the casual agent of sweet potato virus disease in Italy. Plant Pathol 55:818 Perez-Egusquiza Z, Ward LI, Clover GRG, Fletcher JD (2009) Detection of Sweet potato virus 2 in sweet potato in New Zealand. Plant Dis 93:427 Pozzer L, Dusi AN, Kitajima EW (1993) Aphid transmission of Sweet potato feathery mottle virus. Fitopatol Bras 18:274 Prakash S, Tam Y, Zeidan M, Abu-Ras A, Gaba V (2013) First report of Sweet potato virus C infecting sweet potato in Israel. New Dis Rep 28:4 Prasanth G, Hegde V (2008) Occurrence of Sweet potato feathery mottle virus and Sweet potato leaf curl Georgia virus on sweet potato in India. Plant Dis 92:311 Qiao Q, Zhang ZC, Qin YH, Zhang DS, Tian YT, Wang YJ (2011) First report of Sweet potato chlorotic stunt virus infecting sweet potato in China. Plant Dis 95:356 Qiao QI, Zhang ZC, Zhang DS, Qin YH, Tian YT, Wang YJ (2012) Serological and molecular detection of viruses infecting sweet potato in China. Acta Phytopathol Sin 42:10–16 Qin Y, Zhang Z, Qiao Q, Zhang D, Tian Y, Wang Y (2013a) Molecular variability of Sweet potato chlorotic stunt virus (SPCSV) and five potyviruses infecting sweet potato in China. Arch Virol 158:491–495 Qin Y, Zhang Z, Qiao Q, Zhang D, Tian Y, Wang Y, Wang S (2013b) Complete genome sequences of two Sweet potato chlorotic stunt virus isolates from China. Genome Announc 1:218–313 Qin Y, Zhang Z, Qiao Z, Qiao Q, Zhang D, Tian Y, Wang S (2013c) First report of Sweet potato leaf curl Georgia virus on sweet potato in China. Plant Dis 97:1388 Qin Y-H, Li X-C, Zhang Z-C, Qiao Q, Zhang D-S, Wang Y-J, Tian Y-T, Wang S (2016) First report of sweet potato badnavirus A in China. Plant Dis 100:865 Rannali M, Czekaj V, Jones RAC, Fletcher JD, Mu L, Davis R, Gweyer DI, Coutts B, Valkonen JPT (2008) Molecular genetic characterization of Sweet potato virus G (SPVG) isolates from the areas of the pacific ocean and Southern Africa. Plant Dis 92:1313–1320 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Rodriguez Pardina PE, Bejerman N, Luque AV, Di Feo L (2012) Complete nucleotide sequence of an Argentinean isolate of sweet potato virus G. Virus Genes 45(3):593–595 Rosario K, Marr C, Varsani A, Kraberger S, Stainton D, Moriones E, Polston JE, Breitbart M (2016) Begomovirusassociated satellite DNA diversity captured through vector-enabled metagenomic (VEM) surveys using whiteflies (Aleyrodidae). Viruses 8(2):36 Ryu KH, Kim SJ, Park WM (1998) Nucleotide sequence analysis of the coat protein genes of two korean isolates of sweet potato feathery mottle potyvirus. Arch Virol 143:557–562 Sakai J, Mori M, Morishita T, Tanaka M, Hanada K, Usugi T, Nishiguchi M (1997) Complete nucleotide sequence and genome organization of sweet- potato feathery mottle virus (S strain) genomic RNA: the large coding region of the P 1 gene. Arch Virol 142:1553–1562 Sakai J, Kamisoyama S, Yamasaki S, Hanada K (2009) Characterization of an isolate of the common strain group of sweet potato feathery mottle virus from sweet potato [Ipomoea batatas]in Japan. Jpn J Phytopathol 75:156–163 Sharma P, Sahu AK, Verma RK, Mishra R, Choudhary DK, Gaur RK (2014) Current status of Potyvirus in India. Arch Phytopathol Plant Prot 47:906–918 Sim J, Valverde RA, Clark CA (2000) Whitefly transmission of Sweet potato chlorotic stunt virus. Plant Dis 84:1250 Simmons AM, Ling KS, Harrison HF, Jackson DM (2009) Sweet potato leaf curl virus: efficiency of acquisition, retention and transmission by Bemisia tabaci (Hemiptera: Aleyrodidae). Crop Prot 28:1007–1011 Sossah FL, Appiah AS, Oduro V, Amoatey HM, Owusu GK, Oppong A, Lamptey JNL, Carey EE, Fuentes S (2015) Incidence of sweet potato viruses in the coastal savannah agro-ecological zone of Ghana. J Plant Pathol 97:109–117 Souto ER, Sim J, Chen J, Valverde RA, Clark CA (2003) Properties of strains of sweet potato feathery mottle virus and two newly recognized potyviruses infecting sweet potato in the United States. Plant Dis 87:1226–1232 Stobbs LW, Cerkauskas RF, Reynolds LB, McKeown AW (1991) Occurrence of an aphid-transmissible Sweet potato feathery mottle virus in Ontario, Canada. Plant Dis 75(4):430 Tairo F, Kullaya A, Valkonen JPT (2004) Incidence of viruses infecting sweet potato in Tanzania. Plant Dis 88: 916–920 Tairo F, Mukasa SB, Jones RAC, Kullaya A, Rubaihayo PR, Valkonen JPT (2005) Unravelling the genetic diversity of the three main viruses involved in sweet potato virus disease (SPVD) and its partial implications. Mol Plant Pathol 6:119–211 Tairo F, Jones RAC, Valkonen JPT (2006) Potyvirus Complexes in Sweetpotato: occurrence in Australia, Serological and Molecular Resolution, and Analysis of the Sweet potato virus 2 (SPV2) Component. Plant Dis 90:1120–1128 Trenado HP, Lozano G, Valverde RA, Navas-Castillo J (2007) First report of Sweet potato virus G and Sweet potato virus 2 infecting sweet potato in Spain. Plant Dis 91:1687

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Trenado HP, Orílio AF, Márquez-Martín B, Moriones E, Navas-Castillo J (2011) Sweepoviruses cause disease in sweet potato and related Ipomoea spp.: fulfilling Koch’s postulates for a divergent group in the genus Begomovirus. PLoS One 6(11):e27329 Tugume AK, Mukasa SB, Kalkkinen N, Valkonen JP (2010a) Recombination and selection pressure in the ipomovirus sweet potato mild mottle virus (Potyviridae) in wild species and cultivated sweet potato in the centre of evolution in East Africa. J Gen Virol 91:1092–1108 Tugume AK, Cuéllar WJ, Mukasa SB, Valkonen JP (2010b) Molecular genetic analysis of virus isolates from wild and cultivated plants demonstrates that East Africa is a hotspot for the evolution and diversification of sweet potato feathery mottle virus. Mol Ecol 19(15):3139–3156 Tugume AK, Amayo R, Weinheimer I, Mukasa SB, Rubaihayo PR, Valkonen JP. (2013) Genetic variability and evolutionary implications of RNA silencing suppressor genes in RNA1 of sweet potato chlorotic stunt virus isolates infecting sweet potato and related wild species. PLoS One. https://doi.org/10.1371/journal.pone.0081479 Tzanetakis IE, Martin RR, Wintermantel WM (2013) Epidemiology of Criniviruses: an emerging problem in world agriculture. Front Microbiol 4:119. https://doi.org/10.3389/fmicb.2013.00119 Untiveros M, Fuentes S, Kreuze J (2008) Molecular variability of sweet potato feathery mottle virus and other potyviruses infecting sweet potato in Peru. Arch Virol 153:473–483 Usugi T, Nakano M, Shinkai A, Hayashi T (1991) Three filamentous viruses isolated from sweetpotato in Japan. Ann Phytopathol Soc Jpn 57:512–521 Usugi T, Nakano M, Onuki M, Maoka T, Hayashi T (1994) A new strain of sweet potato feathery mottle virus that causes russet crack on fleshy roots of some Japanese cultivars of sweet potato. Ann Phytopath Soc Jpn 60:545–554 Valverde RA, Moreira MA (2004) Identification of sweet potato (Ipomoea batatas L.) viruses in Costa Rica. Agron Mesoamericana 15:1–7 Valverde RA, Lozano G, Navas-Castillo J, Ramos A, Valdes F (2004a) First report of Sweet potato chlorotic stunt virus and Sweet potato feathery mottle virus infecting sweet potato in Spain. Plant Dis 88:428 Valverde RA, Sim J, Lotrakul P (2004b) Whitefly transmission of sweet potato viruses. Virus Res 100:123–128 Valverde RA, Kokkinos CD, Clark CA (2004c) Sweet leaf curl virus: detection by molecular hybridization. Phytopathology 94:S105 Vetten HJ, Hoyer U, Maiss E, Jelkmann W (1996) Serological detection and discrimination of geographically diverse isolates of Sweet potato sunken vein clostero virus. Phytopathology 11:100 Wambugu FM (1991) In vitro and epidemiological studies of sweet potato Ipomoea batatas (L.) Lam. Virus Diseases in Kenya. PhD Thesis, University of Bath, 271 p Wang LY, Chen KC, Chen TC, Yeh SD (2007) Nucleotide sequence analysis and detection application of the coat protein gene of Sweet potato latent virus isolated from central Taiwan. Plant Pathol Bull 16:141–148 Wang M, Abad J, Fuentes S, Li R (2013a) Complete genome sequence of the original Taiwanese isolate of Sweet potato latent virus and its relationship to other potyviruses infecting sweet potato. Arch Virol 158(10):2189–2192 Wang L-Y, Cheng Y-H, Wang N-Y, Chen K-C, Yeh S-D (2013b) First report of Sweet potato virus G infecting sweet potato in Taiwan. Plant Dis 97:1260 Wang Y-J, Zhang D-S, Zhang Z-C, Wang S, Qiao Q, Qin Y-H, Tian Y-T (2015) First report on Sweet potato symptomless virus 1 (Genus: Mastrevirus, Family: Geminiviridae) in sweet potato in China. Plant Dis 99(7):1042 Wassawa P (2012) Sweet potato viruses in Uganda: identification of a new virus, a mild strain of an old virus and reversion. PhD thesis, University of Greenwich (UK), p 220 Wu L, Liu H, Abad J, French RD, Li R (2018) Structure and genome organization of a novel Fiji strain of Sweet potato vein clearing virus identified by high-throughput sequencing. Genome Announc 6(24):e00462–e00418 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Yamasaki S, Sakai J, Kamisoyama S, Hanada K (2009a) Characterization of an isolate of the common strain group of sweet potato feathery mottle virus from sweet potato in Japan. Jpn J Phytopathol 75:156–163 Yamasaki S, Sakai J, Kamisoyama S, Hanada K (2009b) Identification of Sweet potato virus G isolated from sweet potato in Japan. Jpn J Phytopathol 75(2):102–108 Yamasaki S, Sakai J, Fuji S, Kamisoyama S, Emoto K, Ohshima K, Hanada K (2010) Comparisons among isolates of sweet potato feathery mottle virus using complete genomic RNA sequences. Arch Virol 155(5):795–800 Yun WS, Lee YH, Kim K-H (2002) First report of Sweet potato latent virus and Sweet potato chlorotic stunt virus isolated from sweet potato in Korea. Plant Pathol J 18(3):126–129 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98: 131–133 Zhang SC, Ling KS (2011) Genetic diversity of sweet potato begomoviruses in the United States and identification of a natural recombinant between sweet potato leaf curl virus and sweet potato leaf curl Georgia virus. Arch Virol 156(6):955–968 Zhou X (2013) Advances in understanding begomovirus satellites. Ann Rev Phytopathol 51:357–381

Ipomoea purpurea (Common morning glory)

1271

Ipomoea indica (Blue morning glory) Family: Convolvulaceae

Weed host

Sweet potato leaf curl virus

(SPLCV)

Synonyms Ipomoea yellow vein virus (IYVV) Taxonomic position Genus: Begomovirus

Family: Geminiviridae

SPLCV infection in plants of Ipomoea indica was reported from Greece, Italy, and Spain (Banks et al. 1999; Fiallo-Olive et al. 2014). The virus-infected blue morning glory plants exhibit yellow vein symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci (biotype B) in a circulative non-propagative manner. The virus is not transmissible by mechanical inoculation, nor by contact between plants. The virus is transmissible by grafting. For more details of SPLCV, refer to Ipomoea batatas.

Sweet potato leaf curl Canary virus Taxonomic position Genus: Begomovirus

(SPLCCV)

Family: Geminiviridae

SPLCCV infection in plants of Ipomoea indica was reported from Spain (Lozano et al. 2009). The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of SPLCCV, refer to Ipomoea batatas.

References Banks GK, Bedford ID, Beitia FJ, Rodriguez-Cerezo E, Markham PG (1999) A novel geminivirus of Ipomoea indica (Convolvulaceae) from Southern Spain. Plant Dis 83:486 Fiallo-Olive E, Katis NI, Navas-Castillo (2014) First report of Sweet potato leaf curl virus on blue morning glory in Greece. Plant Dis 98:700 Lozano G, Trenado HP, Valverde RA, Navas-Castillo J (2009) Novel begomovirus species of recombinant nature in sweet potato (Ipomoea batatas) and Ipomoea indica: taxonomic and phylogenetic implications. J Gen Virol 90:2550–2562

Ipomoea purpurea (Common morning glory) Family: Convolvulaceae

Weed host

Sweet potato leaf curl virus Synonyms Ipomoea yellow vein virus (IYVV)

(SPLCV)

I

1272

Taxonomic position Genus: Begomovirus

Ipomoea trichocarpa (Cotton morning glory)

Family: Geminiviridae

SPLCV infection in plants of Ipomoea purpurea was reported from India, the United States, Sicily, and China (Yang et al. 2009; Geetanjali et al. 2013). The virus-infected common morning glory plants exhibit yellow vein and leaf curl symptoms. Generally, association of betasatellites with SPLCV was not known. In India, however, two different betasatellites, Croton yellow vein mosaic betasatellite and Papaya leaf curl betasatellite, were identified with SPLCV in the leaf curl and yellow vein diseases of I. purpurea (Geetanjali et al. 2013). The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The virus is not transmissible by mechanical sap-inoculation, and contact between plants. The virus is transmissible by grafting. For more details of SPLCV, refer to Ipomoea batatas.

References Geetanjali AS, Shilpi S, Mandal B (2013) Natural association of two different betasatellites with Sweet potato leaf curl virus in wild morning glory (Ipomoea purpurea) in India. Virus Genes 47:184–188 Yang CX, Wu ZJ, Xie LH (2009) First report of the occurrence of Sweet potato leaf curl virus in Tall morning glory (Ipomoea purpurea) in China. Plant Dis 93:764

Ipomoea trichocarpa (Cotton morning glory) Family: Convolvulaceae

Weed host

Sweet potato feathery mottle virus Taxonomic position Genus: Potyvirus

(SPFMV)

Family: Potyviridae

SPFMV infection in plants of Ipomoea trichocarpa was reported from Louisiana, USA (Clark et al. 1986). The virus-infected cotton morning glory plants exhibit chlorotic spotting and mild chlorotic vein-banding symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SPFMV, refer to Ipomoea batatas.

References Clark CA, Derrick KS, Pace CS, Watson B (1986) Survey of wild Ipomoea spp. as potential reservoirs of sweet potato feathery mottle virus in Louisiana. Plant Dis 70:931–932

Iris spp. (Iris)

1273

Iresine spp. Family: Amaranthaceae

Ornamental

Iresine viroid 1

(IrVd-1)

Taxonomic position Genus: Pospiviroid

Family: Pospiviroidae

Geographical distribution IrVd-1 was detected from plants of Iresine spp. (Iresine herbstii) in Germany (Spieker 1996). Symptoms and host(s) The viroid-infected iresine plants have not shown any macroscopic symptoms. Since tomato was not the host for IrVd-1 (Spieker 1996), it seems unlikely that IrVd-1 poses a threat to potato and tomato cultivation. IrVd-1 has also been identified in Alteranthera sessilis in India (Singh et al. 2006). Transmission The viroid is transmissible mechanically to I. herbstii (Spieker 1996). Etiology and genome properties The genome contains single-stranded circular RNA of 370 nt (X95734 = NC_003613) (Spieker 1996; Gora-Sochacka 2004; Giguere et al. 2014).

References Giguere T, Raj Adkar-Purushothama C, Perreault J-P (2014) Comprehensive secondary structure elucidation of four genera of the family Pospiviroidae. PLoS One 9(6):e98655. https://doi.org/10.1371/journal.pone.0098655 Gora-Sochacka A (2004) Viroids: unusual small pathogenic RNAs. Acta Biochim Pol 51(3):587–607 Singh RP, Dilworth AD, Baranwal VK, Gupta KN (2006) Detection of Citrus exocortis viroid, Ireseine viroid, and Tomato chlorotic dwarf viroid in new ornamental host plants in India. Plant Dis 90(11):1457 Spieker RL (1996) The molecular structure of Iresine viroid, a new viroid species from Iresine herbstii (‘beefsteak plant’). J Gen Virol 77:2631–2635

Iris spp. (Iris) Family: Iridaceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

Family: Secoviridae

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Iris spp. (Iris)

ArMV infection in plants of Iris hollandica was reported from Lithuania (Samuitiene et al. 2008). The virus-infected iris plants exhibit leaf symptoms as pinpoint spots on tips and a light green mosaic pattern which may be more conspicuous on the flower bud sheaths. The virus is transmitted by the nematode vectors Xiphinema diversicaudatum and X. coxi in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Iris spp. was reported from India, Netherlands, the UK, the USA, the Czech Republic, and Israel (Alper and Loebenstein 1981; Hammond et al. 1985; Hammond and Lawson 1988; Kulshrestha et al. 2006a; Duraisamy and Pokorny 2009). In iris BYMV causes variable foliar mosaic symptoms which may be very intense, especially on the central portion of the plant and also with flower breaking symptoms (Hammond et al. 1985). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Broad bean wilt virus Taxonomic position Genus: Fabavirus

(BBWV)

Family: Secoviridae

BBWV (but without specifying either of the currently recognized species BBWV-1 or BBWV-2) was reported from the United Kingdom in plants of rhizomatous iris (Bailiss et al. 1975). The virus-infected iris plants exhibit chlorotic leaf streaking symptoms. The virus is transmitted by aphid vectors in a nonpersistent manner, and also through mechanical sap-inoculation. It is not clear whether isolates from iris are BBWV-1 or BBWV-2. For more details of BBWV, refer to Vicia faba.

Butterfly flower mosaic virus Taxonomic position Genus: Potyvirus

(BFMV)

Family: Potyviridae

Geographical distribution The virus-infected plants of Iris japonica were reported from China and Japan (Chen et al. 2008). Symptoms and host(s) The virus-infected iris plants exhibit mosaic symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically sap-transmissible.

Iris spp. (Iris)

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Virion properties and genome The virions are non-enveloped, flexuous filaments, 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 9.5–10 kb. Partial genome sequences are available (AM774001; GQ240300) (Wylie et al. 2017).

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Iris spp. was reported from China, Lithuania, the Czech Republic, and the Netherlands (Chen 2003; Samuitiene and Navalinskiene 2008; Duraisamy and Pokorny 2009; Lu et al. 2016). The virus-infected plants exhibit leaf symptoms appearing as a light green mosaic pattern which may be more conspicuous on the flower bud sheaths. Symptoms on flowers are expressed by spots of various colors and shapes. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Iris spp. was reported from the Netherlands (Derks and Lemmers 1996). On iris INSV induces yellowish discoloration of inner leaves or sometimes necrotic spots and stripes. The virus is transmitted by a thrips vector Frankliniella occidentalis in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Iris fulva mosaic virus Taxonomic position Genus: Potyvirus

(IFMV)

Family: Potyviridae

Geographical distribution IFMV infection in plants of Iris spp. was reported from the USA (Travis 1957; Barnett and Alper 1977; Barnett 1991).

Symptoms and host(s) The virus-infected iris plants exhibit faint mosaic symptoms with yellowish streaks on the leaves and dark teardrop-shaped markings on flowers.

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Iris spp. (Iris)

Transmission The virus is transmitted by aphid vectors Myzus persicae, Microsiphum euphorbiae, and Aphis fabae in a non-persistent manner. The virus is mechanically sap-transmissible to less than three plant families. The virus is not seed-transmitted. Virion properties and genome The virions are non-enveloped, flexuous filaments 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9.5–10 kb (Revers and Garcia 2015; Wylie et al. 2017).

Iris germanica leaf stripe virus

(IGLSV)

Taxonomic position IGSLV is a tentative member of the family Rhabdoviridae. Geographical distribution IGLSV infection was first reported in plants of Iris germanica from Spain by Rubio-Huertos (1978). The virus spreads in Spain. Symptoms and host(s) The virus-infected iris plants exhibit conspicuous yellow leaf stripes symptoms. Transmission No vector or transmission information is available. Virion properties and genome The virions are bacilliform or bullet-shaped of c.48–52  320–330 nm. The genome is negative-sense RNA of > or equal to 12 kb.

Iris mild mosaic virus Taxonomic position Genus: Potyvirus

(IMMV)

Family: Potyviridae

Geographical distribution IMMV infection was first identified in plants of Iris spp. in the Netherlands (Van Slogteren 1958). The virus is probably distributed worldwide (Derks et al. 1985; Kulshrestha et al. 2006b; Pham et al. 2011; Nateqi et al. 2017). Symptoms and host(s) Leaf symptoms appear as a faint mosaic which may be more conspicuous on the flower bud sheath. Often the top halves of the leaves are light green to yellow green with small dark green stripes. Flower symptoms are limited to a few small dark flecks in the falls (the lower petals) (Hammond et al. 1985). Mild mosaic symptoms are observed on different iris species like bulbous irises, Iris xiphiodes (English iris), I. xiphium (Spanish iris), I. tingitana x I. xiphium (Dutch iris), I. reticulata, and I. danfordiae (Asjes 1979).

Iris spp. (Iris)

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Transmission The virus is transmitted by aphid vectors such as Aphis gossypii, Macrosiphum euphorbiae, and Myzus persicae, in a non-persistent manner but is not by Macrosiphoniella sanborni. The virus is mechanically sap-transmissible. The virus is not transmitted by seed (Derks et al. 1985). Virion properties and genome The virions are non-enveloped, flexuous filaments, with a clear modal length of 700 nm long and 12 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA (Revers and Garcia 2015). Partial genome sequences are available (DQ436918, JF320812; JN127338) (Wylie et al. 2017).

Iris severe mosaic virus

(ISMV)

Synonyms Bearded iris mosaic virus; Beardless iris mosaic virus Taxonomic position Genus: Potyvirus

I Family: Potyviridae

Geographical distribution ISMV infection in plants of Iris spp. was reported from Australia, China, the former Czechoslovakia, Denmark, Israel, Iran, Japan, the Netherlands, Sweden, the United Kingdom, and the USA (and probably wherever bulbous and rhizomatous irises and crocus are grown) (Brierley and Smith 1948; Travis 1957; Barnett et al. 1971; Van der Vlugt et al. 1993; Kulshrestha et al. 2004; Yan et al. 2010; Nateqi et al. 2015). Symptoms and host(s) ISMV-infected iris plants exhibit symptoms vary from mild to severe; mild symptoms which consist of yellowish green stripes in the outer leaves or on developing middle leaves. Severely affected plants show wide, pale green, and yellowish-green stripes. The entire plant may show various degrees of stunting, from slight stunting to severe dwarfing, which results in a short flower stalk for cut blooms. The quality of the flower is also inferior due to dark teardrop markings (color breaking) in diseased standards or falls, or both, of white, blue, and lavender cultivars. Flowers may be reduced in size and are often twisted to one side. Bud sheaths may show bluish green blotches on a pale green background or, less commonly, yellowish streaks. The most intense mottling symptoms appear on the bud sheaths and youngest leaves and stems under cool conditions (Hammond et al. 1985). Symptoms are typically less severe in rhizomatous iris, with faint light green mosaic or occasional light green or yellow mosaic (Barnett et al. 1971). Transmission The virus is transmitted by aphid vectors Macrosiphum euphorbiae and Myzus persicae in a nonpersistent manner. The virus is also transmissible by mechanical sap-inoculation. The virus is not transmissible by grafting and also not by contact between plants. Virion properties and genome The virions are non-enveloped, filaments usually flexuous, with a clear modal length of 750 nm and 12 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 10,423 nt

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Iris spp. (Iris)

(KT692938 = NC_029076) (Van der Vlugt et al. 1994; Kulshrestha et al. 2004; Revers and Garcia 2015; Li et al. 2016; Wylie et al. 2017).

[doc.name]}>Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

Geographical distribution IYSV was reported in plants of Iris spp. from the Netherlands, the Czech Republic, Iran, and the USA (Derks and Lemmers 1996; Bag et al. 2009; Duraisamy and Pokorny 2009; Rafizadeh et al. 2013; Basavaraj et al. 2017). Symptoms and host(s) The virus-infected iris showed yellow and sometimes necrotic spots on leaves (Derks and Lemmers 1996). Symptoms were later reported to consist of chlorotic spots that developed into yellow and necrotic spots (Cortes et al. 1998). The natural host range of this virus includes various weeds, but is mainly confined to allium species, and it is continuously being reported from different allium-growing regions, compared to other tospoviruses reported from various vegetable and oilseed crops of economic importance. Transmission The virus is transmitted by thrips vectors Thrips tabaci and Frankliniella fusca in a persistentpropagative manner. The virus is also mechanically sap-transmissible (Srinivasan et al. 2012). Virion properties and genome The virions are spherical and membrane-bound 80–120 nm in diameter. The genome comprises three unique molecules of negative or ambisense ssRNA, designated L (large) 8880 nt (FJ623474 = NC_029799), M (medium) 4821 nt (AF214014 = NC_029798), and S (small) 3105 nt (AF001387 = NC_029800) (Cortes et al. 1998; Bag et al. 2009, 2010).

Japanese iris necrotic ring virus Taxonomic position Genus: Betacarmovirus

(JINRV)

Family: Tombusviridae

Geographical distribution JINRV infection was reported in plants of Iris kaempferi in Japan and Australia (Yasukawa et al. 1982, 1991; Wylie et al. 2012). Symptoms and host(s) The infected iris plants exhibit symptoms of spindle-shaped necrotic streaks or rings in leaves. Transmission The virus is mechanically sap-transmissible to fewer than three plant families. Aphids failed to transmit this virus disease.

Iris spp. (Iris)

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Virion properties and genome The virions are isometric 30 nm in diameter and have T = 3 icosahedral symmetry. The genome consists of a single molecule of linear positive-sense ssRNA of 4014 nt (D86123 = NC_002187) (Takemoto et al. 2000).

Narcissus latent virus

(NLV)

Synonyms Iris mild yellow mosaic virus (IMYMV)

Taxonomic position Genus: Macluravirus

Family: Potyviridae

NLV infection in plants of Iris spp. was reported from Australia, Germany, the Netherlands, the United Kingdom and the USA (Asjes 1979; Brunt 1977; Derks et al. 1985; Hammond et al. 1985; Wei et al. 2007). The virus-infected leaves of iris show a yellow-green mosaic pattern, especially at the margins and tips, which is typically more prominent soon after emergence than later in the growing season. Necrotic spots appear on the top half of the leaves, mostly after flowering (Hammond et al. 1985). The virus is transmitted by aphid vectors Aphis gossypii, Acyrthosiphon pisum, and Myzus persicae in a non-persistent manner, and also by mechanical sap-inoculation. For more details of NLV, refer to Narcissus spp.

Narcissus mosaic virus Taxonomic position Genus: Potexvirus

(NMV)

Family: Alphaflexiviridae

NMV was reported in plants of Iris xiphium for the first time in Australia (Wylie et al. 2014). The virusinfected iris plants exhibit chlorotic streaks on the leaves and spathe symptoms. The virus is mechanically sap-transmissible. For more details of NMV, refer to Narcissus spp.

Ornithogalum mosaic virus Taxonomic position Genus: Potyvirus

(OrMV)

Family: Potyviridae

OrMV infection in plants of bulbous iris and Iris tingitana was reported from New Zealand (Wei et al. 2006); in bulbous iris in India exhibiting mottle-mosaic, chlorotic spots on leaves and deformed flowers (Chandel et al. 2006); and in bulbous iris in Korea shows severe mosaic (Yoon and Ryu 2002). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of OrMV, refer to Ornithogalum spp.

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Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

Iris spp. (Iris)

(TMV)

Family: Virgaviridae

TMV was reported in plants of bulbous iris (Iris hollandica) from the Netherlands and Lithuania (Asjes 1979; Samuitiene et al. 2011). The virus-infected iris plants exhibit chlorotic mottle progressing to necrotic mottling and streaking of leaves. There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV has occasionally been detected, but not associated with particular symptoms, in bulbous iris (Iris hollandica) in the Netherlands (Asjes 1979), and also reported in the Czech Republic (Duraisamy and Pokorny 2009). TRV is also occasionally detected in rhizomatous irises in the USA. The virus is transmitted by nematode vectors and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV was reported in plants of Iris spp. from the Netherlands and the United Kingdom (Travis and Brierley 1957; Brunt 1975; Asjes 1979; Samuitiene et al. 2011). The virus-infected iris plants exhibit chlorotic streaking and ring-spot symptoms. The virus is transmitted by nematode vectors in a nonpersistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Iris spp. was reported from Lithuania (Samuitiene et al. 2011). The virusinfected iris plants exhibit chlorotic and ringspots on leaves. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ToRSV, refer to Solanum lycopersicum.

Iris spp. (Iris)

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Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Iris spp. was reported from the Netherlands (Yamamoto and Ohata 1977; Derks and Lemmers 1996; Adkins et al. 2003). TSWV on iris induces yellowish discoloration of inner leaves and sometimes necrotic spots and stripes. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato zonate spot virus

(TZSV)

Taxonomic position TZSV is a tentative member of the genus Orthotospovirus and family Tospoviridae TZSV infection in plants of Iris tectorum was reported from China (Liu et al. 2015). The virus-infected iris plants exhibit symptoms of chlorotic spots on younger leaves and necrosis on older leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner. For more details of TZSV, refer to Solanum lycopersicum.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV has been reported from plants of bulbous iris in Japan (Inouye and Mitsuhata 1978). The virusinfected iris plants exhibit leaf necrosis and mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also through mechanical sap-inoculation. For more details of TuMV, refer to Brassica rapa.

References Adkins S, Breman L, Baker CA, Wilson S (2003) First report of Tomato spotted wilt virus in Blackberry Lily in North America. Plant Dis 87:102 Alper M, Loebenstein G (1981) Bean yellow mosaic virus in bulbous irises in Israel. Plant Dis 65:694–695 Asjes CJ (1979) Viruses and virus diseases in Dutch bulbous irises (Iris hollandica) in the Netherlands. Neth J Plant Pathol 85:269–279 Bag S, Druffel KL, Salewsky T, Pappu HR (2009) Nucleotide sequence and genome organization of the medium RNA of IYSV (genus Tospovirus, family Bunyaviridae) from the Unites States. Arch Virol 154:715–718 Bag S, Druffel KL, Pappu HR (2010) Structure and genome organization of the large RNA of iris yellow spot virus (genus Tospovirus, family Bunyaviridae). Arch Virol 155:275–279 Bailiss KW, Brunt AA, Dale WT (1975) The natural occurrence of broad bean wilt virus in rhizomatous iris and spinach. Plant Pathol 24:60–61 Barnett OW (1991) Iris fulva mosaic virus, AAB Descriptions of Plant Viruses No. 310. Association of Applied Biologists, Wellesbourne

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Iris spp. (Iris)

Barnett OW, Alper M (1977) Characterization of Iris fulva mosaic virus. Phytopathology 67:448–454 Barnett OW, de Zoeten GA, Gaard G (1971) Bearded iris mosaic virus: transmission, purification, inclusions, and its differentiation from Bulbous iris mosaic. Phytopathology 61:926–932 Basavaraj, Mandal B, Gawande SJ, Renukadevi P, Holkar SK, Krishnareddy M, Ravi KS, Jain RK (2017) The occurrence, biology, serology and molecular biology of Tospoviruses in Indian agriculture. In: Mandal B, Rao GP, Baranwal VK, Jain RK (eds) A century of plant virology in India. Springer Nature, Singapore, pp 445–474. ISBN 978-981-10-5671-0 Brierley P, Smith FE (1948) Two additional mosaic diseases of iris. Phytopathology 38:575–575 Brunt AA (1975) Viruses and virus diseases of irises in Britain. Acta Hortic 47:45–50 Brunt AA (1977) Some hosts and properties of Narcissus latent virus, a carlavirus commonly infecting narcissus and bulbous iris. Ann Appl Biol 87:355–364 Chandel V, Kulshrestha S, Hallan V, Zaidi AA (2006) Natural infection of Ornithogalum mosaic virus on Iris from India. Plant Pathol 55:284 Chen YK (2003) Occurrence of cucumber mosaic virus in ornamental plants and perspectives of transgenic control. Ph.D. Thesis, Wageningen University, The Netherlands. p 144 Chen J, Shi YH, Li MY, Adams MJ, Chen JP (2008) A new potyvirus from butterfly flower (Iris japonica Thunb.) in Zhejiang, China. Arch Virol 153:567–569 Cortes I, Livieratos IC, Derks A, Peters D, Kormelink R (1998) Molecular and serological characterization of Iris yellow spot virus, a new and distinct tospovirus species. Phytopathology 88:1276–1282 Derks AFLM, Lemmers MEC (1996) Detection of tospoviruses in bulbous crops and transmissibility by vegetative propagation. Acta Hortic 432:132–137 Derks AFLM, Hollinger THC, Vinkvan den Abeele JL (1985) Identification and symptom expression of four elongated viruses infecting bulbous irises. Acta Hortic 164:309–318 Duraisamy GS, Pokorny R (2009) Survey of virus pathogens in gladiolus, iris and tulips in the Czech Republic. Acta Univ Agric Silviculturae Mendelianae Brun 57:79–86 Hammond J, Lawson RH (1988) An improved purification procedure for preparing potyviruses and cytoplasmic inclusions from the same tissue. J Virol Meth 20(3):203–217 Hammond J, Brunt AA, Derks AFLM, Inouye N, Barnett OW, Allen TC, Lawson RH (1985) Viruses infecting bulbous iris: a classification of nomenclature. Acta Hortic 164:395–397 Inouye N, Mitsuhata K (1978) Turnip mosaic virus isolated from iris. Nogaku Kenkyu 57:1–16 Kulshrestha S, Hallan V, Raikhy G, Kumar A, Ram R, Garg ID, Zaidi AA (2004) Molecular characterization of an Iris severe mosaic virus isolate from India. Acta Virol 48:65–67 Kulshrestha S, Hallan V, Raikhy G, Ram R, Zaidi AA, Garg ID (2006a) Incidence of Bean yellow mosaic virus in Iris. Acta Hortic 722:235–240 Kulshrestha S, Hallan V, Raikhy G, Ram R, Garg ID, Haq QMR, Zaidi AA (2006b) Occurrence of Iris mild mosaic potyvirus in cultivated iris in India. Indian J Biotechnol 5:94–98 Li Y, Deng C, Shang Q, Zhao X, Liu X, Zhou Q (2016) The first complete genome sequence of iris severe mosaic virus. Arch Virol 161(4):1069–1072 Liu Y, Huang CJ, Tao XR, Yu HQ (2015) First report of Tomato zonate spot virus in Iris tectorum in China. Plant Dis 99:164 Lu XB, Tang W, Xu XH, Sun HW, Li F, Gao R, Yang SK (2016) First report of Cucumber mosaic virus on Iris tectorum in China. Plant Dis 100:1512 Nateqi M, Koohi Habibi M, Dizadji A, Parizad S (2015) Detection and molecular characterization of the Iris severe mosaic virus-Ir isolate from Iran. J Plant Protect Res 55:235–240 Nateqi M, Dizadji A, Koohi Habibi M, Movi S (2017) First report of Iris mild mosaic virus from Iris xiphium in Iran. J Plant Pathol 99:287–304 Pham KTK, de Kock MJD, Lemmers MEC, Derks AFLM (2011) Molecular identification of potyviruses infecting bulbous ornamentals by the analysis of coat protein (CP) sequences. Acta Hortic 901:167–172 Rafizadeh N, Jafarpour B, Falahati Rastegar M (2013) Detection of Iris yellow spot virus (IYSV) in onion and some of ornamental plants by ELISA and RT-PCR methods in Khorosan Razavi provinces. J Plant Prot (Agric Sci Technol) 27:149–158 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Rubio-Huertos M (1978) A rhabdovirus in Iris germanica. Phytopathol Z 92:294–297 Samuitiene M, Navalinskiene M (2008) Occurrence of Cucumber mosaic cucumovirus on ornamental plants in Lithuania. Zemdirbyste-Agriculture 95:135–143 Samuitiene M, Navalinskiene M, Jackeviciene E (2008) Arabis mosaic virus on ornamental plants. Biologia 54:264–268 Samuitiene M, Navalinskien_e M, Zitikait_e I, Kompanets T, Korotyeyeva A, Bysov A (2011) Detection and identification of viruses affecting irises in Lithuania and Ukraine. Biologiya 59:18–22 Srinivasan R, Sundaraj S, Pappu HR, Diffie S, Riley D, Gitaitis R (2012) Transmission of Iris yellow spot virus by Frankliniella fusca and Thrips tabaci (Thysanoptera: Thripidae). J Ecol Entomol 105:40–47

Isotoma axillaris (Rock isotome)

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Takemoto Y, Kanehira T, Shinohara M, Yamashita S, Hibi T (2000) The nucleotide sequence and genome organization of Japanese iris necrotic ring virus, a new species in the genus Carmovirus. Arch Virol 145:651–657 Travis RV (1957) Virus diseases of iris. Phytopathology 47:454 Travis RV, Brierley P (1957) Tobacco ring spot virus from iris and Easter lily. Plant Dis Reptr 41:254 Van der Vlugt CIM, Derks AFLM, Boonekamp PM, Goldbach RW (1993) Improved detection of Iris severe mosaic virus in secondarily-infected iris bulbs. Ann Appl Biol 122:279–288 van der Vlugt CI, Langeveld SA, Goldbach RW (1994) Molecular cloning and sequence analysis of the 30 -terminal region of iris severe mosaic virus RNA. Arch Virol 136(3–4):397–406 Van Slogteren DHM (1958) Versl. Werkzaamh. Bloemboll Lisse, 1957 Wei T, Pearson MN, Cohen D (2006) First report of Ornithogalum mosaic virus and Ornithogalum virus 2 in New Zealand. Plant Pathol 55:820 Wei T, Pearson MN, Cohen D (2007) First report of Narcissus latent virus in New Zealand. Plant Pathol 56:720 Wylie SJ, Li H, Jones MGK (2012) First report of an isolate of Japanese iris necrotic ring virus from Australia. Australas Plant Dis Notes 7:107–110 Wylie SJ, Li H, Liu J, Jones MGK (2014) First report of Narcissus mosaic virus from Australia and from Iris. Australas Plant Dis Notes 9:134 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Yamamoto T, Ohata K (1977) Some properties and electron microscopy of Tomato spotted wilt virus isolated from blackberry lily (Belamcanda chinensis DC.). Bull Shikoku Agric Exp Sta 30:39–47 Yan S, Qin Z, Jin L, Chen J (2010) A new isolate of Iris severe mosaic virus causing yellow mosaic in Iris ensata Thunb. J Nanosci Nanotechnol 10:726–730 Yasukawa K, Ohki ST, Osaki T, Inouye T (1982) Japanese iris necrotic ring disease, a newly recognized disease caused by Japanese iris necrotic ring virus. Ann Phytopathol Soc Jpn 48:113–114 Yasukawa K, Osaki T, Inouye T (1991) Necrotic ring disease of Japanese Iris (Iris kaempferi Sieb.), a new disease caused by Japanese iris necrotic ring virus. Bull Univ Osaka Pref Ser B Agric Biol 43:21–28 Yoon H-I, Ryu K-H (2002) Molecular identification and sequence analysis of coat protein gene of Ornithogalum mosaic virus isolated from Iris plant. Plant Pathol J 18:251–258

Isotoma axillaris (Rock isotome) Family: Campanulaceae

Ornamental

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Isotoma axillaris was reported from Italy (Bellardi et al. 2011). The virusinfected rock isotome plants exhibit symptoms of small necrotic concentric rings and necrosis of the leaf lamina. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

References Bellardi MG, Cavicchi L, Casadei P, Vicchi V, Bozzano G (2011) First report of Impatiens necrotic spot virus infecting Isotoma axillaris. J Plant Pathol 93:S4.26

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Ixia spp.

Ixia spp. Family: Iridaceae

Ornamental

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Ixia hybrida was reported from Japan and the Netherlands (Bellardi et al. 1987; Tsuji et al. 1996). The virus-infected ixia plants exhibit mild mottle or mosaic on the leaves and color break on the flowers but sometimes had no visible symptoms. The virus is transmitted by an aphid vector, Myzus persicae, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

References Bellardi MG, Pisi A, Masenga V (1987) Bean yellow mosaic virus infecting Ixia bulbs imported from Holland. Phytopathol Mediterr 26:165 Tsuji T, Maeda T, Kondo H, Inouye N (1996) Characterization of Bean yellow mosaic virus from Ixia hybrida. Bull Res Inst Bioresour Okayama Univ 4:201–213

Ixora spp. Family: Rubiaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Ixora spp. was reported from the USA and the Philippines (Waterworth and Povish 1975; Moriones et al. 1991; McGarvey et al. 1995). The virus-infected ixora plants exhibit conspicuous mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Tomato mosaic virus Taxonomic position Genus: Tobamovirus

(ToMV)

Family: Virgaviridae

Ixora spp.

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Plants of Ixora duffii “Super King” showing foliar mosaic observed in Taiwan and were shown to be infected with ToMV (Tsai et al. 2008). No vector is involved in the spread of this virus. This virus is transmissible by mechanical sap-inoculation, by grafting, and by contact between plants. For more details of ToMV, refer to Solanum lycopersicum.

References McGarvey P, Tousignant M, Geletka L, Cellini F, Kaper JM (1995) The complete sequence of a Cucumber mosaic virus from Ixora that is deficient in replication of satellite RNAs. J Gen Virol 76:2257–2270 Moriones E, Fraile A, García-Arenal F (1991) Host-associated selection of sequence variants from a satellite RNA of Cucumber mosaic virus. Virology 184:465–468 Tsai JC, Chen TH, Chang CA (2008) Identification and characterization of a Tomato virus isolate infecting Ixora (Ixora duffii cv. ‘Super King’). Plant Pathol Bull 17:143–155 Waterworth HE, Povish WR (1975) A virus related to Cucumber mosaic virus isolated from imported Ixora plants. Phytopathology 65:728–729

I

J

Jacquemontia spp. Family: Convolvulaceae

Ornamental

Jacquemontia mosaic Yucatan virus Taxonomic position Genus: Begomovirus

(JacMYuV)

Family: Geminiviridae

Geographical distribution JacMYuV infection in plants of Jacquemontia pentantha was reported from Yucatan (Mexico) (TorresHerrera et al. unpublished - JQ821386). Symptoms and host(s) The virus-infected jacquemontia plants exhibit foliar yellow mosaic symptoms. Transmission The transmission of JacMYuV has not been investigated. It is likely that, in common with other begomoviruses, JacMYuV will be transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. Similarly it is likely that the virus will not to be either mechanically or seed-transmitted. Virion properties and genome The structure of the virions of JacMYuV has not been investigated. In common with all geminiviruses, the virions of JacMYuV are likely geminate (twinned icosahedra). JacMYuV is typical of bipartite begomoviruses native to the New World. The genome consists of two circular, single-stranded DNA components. DNA-A consists of 2602 nt (JQ821386) and DNA-B of 2591 nt (JQ821387) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). Typical of the majority of bipartite begomoviruses native to the New World, the DNA-A component of JacMYuV encodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for JacMYuV. © Springer Nature India Private Limited 2019 K. S. Sastry et al., Encyclopedia of Plant Viruses and Viroids, https://doi.org/10.1007/978-81-322-3912-3

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Jacquemontia spp.

Jacquemontia yellow mosaic virus Taxonomic position Genus: Begomovirus

(JacYMV)

Family: Geminiviridae

Geographical distribution JacYMV infection in plants of Jacquemontia spp. was reported from Venezuela (Fiallo-Olive et al. 2014). Symptoms and host(s) The virus-infected jacquemontia plants exhibit foliar yellow mosaic symptoms. Transmission The transmission of JacYMV has not been investigated. It is likely that, in common with other begomoviruses, JacYMV will be transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. Similarly it is likely that the virus will not to be either mechanically or seed-transmitted. Virion properties and genome The structure of the virions of JacYMV has not been investigated. In common with all geminiviruses, the virions of JacYMV are likely geminate (twinned icosahedra). JacYMV is typical of bipartite begomoviruses native to the New World. The genome consists of two circular, single-stranded DNA components. DNA-A consists of 2611 nt (KF661331) and DNA-B of 2578 nt (KF661332) (Briddon 2001; Fiallo-Olive et al. 2014; Brown et al. 2015; Zerbini et al. 2017). Typical of the majority of bipartite begomoviruses native to the New World, the DNA-A component of JacYMV encodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for JacYMV.

Jacquemontia yellow vein virus Taxonomic position Genus: Begomovirus

(JacYVV)

Family: Geminiviridae

Geographical distribution JacYVV infection in plants of Jacquemontia tamnifolia was reported from Venezuela (Fiallo-Olive et al. 2017). Symptoms and natural host(s) The virus-infected jacquemontia plants exhibit foliar yellow vein symptoms. Transmission The transmission of JacYVV has not been investigated. It is likely that, in common with other begomoviruses, JacYVV will be transmitted by the whitefly Bemisia tabaci in a circulative,

Jacquemontia spp.

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non-propagative manner. Similarly it is likely that the virus will not to be either mechanically or seedtransmitted. Particle morphology The structure of the virions of JacYVV has not been investigated. In common with all geminiviruses, the virions of JacYVV are likely geminate (twinned icosahedra). JacYVV is typical of bipartite begomoviruses native to the New World. The genome consists of two circular, single-stranded DNA components. DNA-A consists of 2585 nt (KY617094, KY624376) and DNA-B of 2543 nt (KY617095, KY617096) (Brown et al. 2015; Fiallo-Olive et al. 2017; Zerbini et al. 2017). Typical of the majority of bipartite begomoviruses native to the New World, the DNA-A component of JacYVV encodes five genes, one in the virion-sense and four in the complementarysense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for JacYVV.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Jacquemontia tamnifolia was reported from the USA (Schuster and Halliwell 1994; Mullis et al. 2009). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer Index of Viruses. Springer New York, p 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Fiallo-Olive E, Chirinos DT, Geraud-Pouey F, Moriones E, Navas-Castillo J (2014) Complete genome sequence of Jacquemontia yellow mosaic virus, a novel begomovirus from Venezuela related to other New World bipartite begomoviruses infecting Convolvulaceae. Arch Virol 159:1857–1860 Fiallo-Olive E, Chirinos DT, Geraud-Pouey F, Navas-Castillo J (2017) Complete genome sequence of jacquemontia yellow vein virus, a novel begomovirus infecting Jacquemontia tamnifolia in Venezuela. Arch Virol 162(8):2463–2466 Mullis S, Bertrand PF, Brown SL, Csinos A, Diaz-Perez JC, Gitaitis R, Hickman LL, Johnson A, LaHue SS, Martinez N, McPherson RM, Nischwitz C, Reay-Jones F, Riley D, Sherwood J, Stevenson K, Wells L (2009) Tospoviruses in Solanaceae and other crops in the coastal plain of Georgia. University of Georgia, College of Agricultural and Environmental Sciences, Bulletin 1354, pp 51 Schuster GL, Halliwell RS (1994) Six new hosts of tomato spotted wilt virus in Texas. Plant Dis 78:100 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98: 131–133

J

1290

Jasminum spp.

Jasminum spp. Family: Oleaceae

Ornamental

(AEV)

Ageratum enation virus

Synonyms Jasminum sambac leaf curl virus Lakshmangarh Taxonomic position Genus: Begomovirus

Family: Geminiviridae

AEV, as ‘Jasminum sambac leaf curl virus Lakshmangarh’ (JN998445) caused leaf curl symptoms on Jasminum sambac in India (Marwal et al. 2012, 2013). The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of AEV, refer to Ageratum spp.

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

(AMV)

Family: Bromoviridae

AMV was reported from plants of Jasminum multiflorum in the USA (Waterworth 1971). The virus-infected Jasminum plants exhibited mild mosaic symptoms. The virus is transmitted by a large number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

ArMV was reported in the United Kingdom (Cooper and Sweet 1976) and in the USA in plants of Jasminum officinale and J. mesnyi (= J. primulinum) imported from Scotland (Waterworth 1975). The virus-infected jasminum plants exhibited symptoms of yellow rings and blotches. This virus is transmitted by nematode vectors (Xiphinema spp.) in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Cotton leaf curl Kokhran virus Taxonomic position Genus: Begomovirus

(CLCuKoV)

Family: Geminiviridae

Jasminum spp.

1291

CLCuKoV infection in plants of Jasminum sambac was reported from Pakistan (Akram et al. 2017). The virus-infected jasminum plants exhibit mild leaf curling, yellowing, and vein thickening symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci, in a circulative non-propagative manner. The virus is not transmissible by mechanical inoculation, nor by contact between plants. For more details of CLCuKoV, refer to Gossypium spp.

(CMV)

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV was reported in plants of Jasminum spp. in Denmark and the USA (Waterworth 1971; Anon. 1982; Usharani et al. 2016). The virus-infected jasminum plants exhibited chlorotic ringspots and irregular chlorotic patches. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Groundnut bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(GBNV)

Family: Tospoviridae

GBNV infection in plants of Jasminum sambac was reported from India (Sujitha et al. 2013). The virus-infected jasmine plants showed mosaic and chlorotic spots on the young leaves. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of GBNV, refer to Arachis hypogaea.

Jasmine virus C

(JaVC)

Taxonomic position JaVC is a tentative member of the genus Carlavirus and the family Betaflexiviridae. Geographical distribution JaVC infection in plants of Jasminum sambac was reported from Italy and Taiwan (Bellardi and Bertaccini 1991; Lin 2012). Symptoms and host(s) The virus-infected jasminum plants exhibit foliar yellow mosaic symptoms. Transmission The virus is presumed to be transmitted by aphids. Virion properties and genome Slightly flexuous particles of 600–630 nm were observed in sap extracts; the genome consists of a single molecule of single-stranded positive-sense RNA of 8506 nt (KX364696 = NC_030926).

J

1292

Jasminum spp.

(JVT)

Jasmine virus T

Synonyms Jasmine ringspot virus; Jasmine yellow mosaic virus (JaYMV) Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution JVT infection in plants of Jasminum sambac was reported from Taiwan, India, and China (Lin et al. 2004; Kaur et al. 2013; Sudheera et al. 2014; Tang et al. 2016). Symptoms and host(s) The virus-infected jasminum plants exhibit vein-clearing, followed by irregular chlorotic spots and severe mosaic symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, 720–750 nm long, and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9590 nt (KT222674 = NC_029051) (Tang et al. 2016; Wylie et al. 2017).

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Jasminum mesnyi was reported from the USA and the UK (Morton et al. 1977). The virus-infected jasminum plants produce fewer flowers, shorter internodes, and weaker roots than healthy plants, and more than half of the infected plants die. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Jasminum sambac was reported from India (Seshadri Goud et al. 2013). The virus-infected jasminum plants exhibit symptoms of wilting and necrotic streak of the petioles and branches, with proliferation of axillary shoots having small leaves. The disease was widespread and caused significant yield loss. The virus is transmitted by the thrips vectors, the virus present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is mechanically

Jasminum spp.

1293

sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

(ToMV)

Tomato mosaic virus Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

ToMV infection in plants of Jasminum multiflorum and J. gracile was reported from the USA (Kamenova et al. 2006; Fillmer et al. 2015). The virus-infected jasminum plants exhibit symptoms of mottling, chlorotic ringspots, and chlorotic line patterns. No vector is involved in the spread of this virus. This virus is transmissible by mechanical sap-inoculation, by grafting, and by contact between plants. For more details of ToMV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

J Family: Tospoviridae

TSWV infection in plants of Jasminum spp. was reported from Iran and Portugal (Louro 1996; Parrella et al. 2003; Ghotbi et al. 2005). The virus-infected jasminum plants were symptomatic; no specific symptoms were described. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow ring virus

(TYRV)

Synonyms Tomato varamin virus (ToVV) Taxonomic position TYRV is a tentative member of the genus Orthotospovirus and family Tospoviridae ToVV infection in plants of Jasminum spp. was reported from Iran (Ghotbi et al. 2005). The virusinfected jasminum plants were symptomatic; no specific symptoms were described. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also through mechanical sap-inoculation. For more details of TYRV, refer to Solanum lycopersicum.

References Akram A, Rasool G, Rehman A, Mansoor S, Briddon RW, Saeed M (2017) Identification of Cotton leaf curl Kokhran virus and multiple satellite molecules infecting Jasminum sambac in Pakistan. J Plant Pathol 99:804 Anon (1982) Annual report, research centre for plant protection. Plant disease and pests in Denmark, 1981. Lynghy, Denmark, p 76

1294

Jatropha spp.

Bellardi MG, Bertaccini A (1991) Virus diseases of ornamental shrubs. VI. Chlorotic spots and variegation of Jasminum sambac. Phytopathol Mediterr 30:67–71 Cooper JI, Sweet JB (1976) The detection of viruses with nematode vectors in six woody hosts. Forestry 49:73–78 Fillmer K, Adkins S, Pongam P, D’Elia T (2015) Complete genome sequence of tomato mosaic virus isolated from jasmine in the United States. Genome Announc 3(4):e00706–e00715 Ghotbi T, Sharaeen N, Winter S (2005) Occurrence of tospoviruses in ornamental and weed species in Markazi and Tehran provinces in Iran. Plant Dis 89:425–429 Kamenova I, Adkins S, Achor D (2006) Identification of Tomato mosaic virus infection in jasmine. Acta Hortic 722:277–283 Kaur C, Kumar S, Snehi SK, Raj SK (2013) Molecular detection of Jasmine potyvirus associated with yellow mosaic symptoms on Jasminum sambac L. in India. Arch Phytopathol Plant Protect 46:1102–1107 Lin YC (2012) Identification of a newly recognized carlavirus and its etiology of yellow mottling symptoms on jasmine. M.S. Thesis, ChaoYang University of Technology, Taiwan Lin YY, Chen TH, Chang CA (2004) Characterization of a new potyvirus isolated from jasmine (Jasminum sambac (L.) Ait) in Taiwan. Plant Pathol Bull 13:69–84 Louro D (1996) Detection and identification of Tomato spotted wilt virus and Impatiens necrotic spot virus in Portugal. Acta Hortic 431:99–105 Marwal A, Prajapat R, Sahu A, Gaur RK (2012) Computational characterization of Begomovirus infecting two ornamental plants: Jasminum sambac and Millingtonia hortensis. Asian J Biol Sci 5:240–249 Marwal A, Sahu A, Prajapat R, Gaur RK (2013) First report of begomovirus infecting two ornamental plants: Jasminum sambac and Millingtonia hortensis. Indian Phytopathol 66:115–116 Morton CS, Barnett OW, Baxter LW (1977) Damage caused by tobacco ringspot virus to poet’s jasmine (Jasminum officinale). Proc. Southern Nurserymen’s Assoc. Res. Cong., 22nd Ann. Rep. 111–114 Parrella G, Gognalons P, Gebre-Selassie K, Vovlas C, Marchoux G (2003) An update of the host range of Tomato spotted wilt virus. J Plant Pathol 85:227–264 Seshadri Goud TE, Vemana K, Reddy DL, Mahammed Khureshee CS, Padma JG, Shabbir S, Venkateswarlu NC, Naik KSS, Sampath Kumar D, Anthony Johnson AM, Subramanyam K (2013) First report of Tobacco streak ilarvirus infecting jasmine and horse gram. New Dis Rep 28:7 Sudheera Y, Vishnu Vardhan GP, Hema M, Krishna Reddy M, Sreenivasalu P (2014) Characterization of a potyvirus associated with yellow mosaic disease of jasmine (Jasminum sambac L.) in Andhra Pradesh, India. Virus Dis 25:394–397 Sujitha A, Bhaskara Reddy BV, Sivaprasad Y, Usha R, Giridhara Krishna T, Sai Gopal DVR (2013) First report of Groundnut bud necrosis virus in jasmine. J Plant Pathol 95:667 Tang Y, Gao F, Yang Z, Wu Z, Yang L (2016) Complete genome analysis of Jasmine virus T from Jasminum sambac in China. Arch Virol 161:2033–2036 Usharani TR, Jalali S, Manasa M, Samuel DK, Krishnareddy M (2016) Identification of mixed infection caused by Badnavirus and CMV in Jasmin (Jasminum multiflorum Roth). In: IVS International Conference on Global Perspectives in Virus Disease Management. VIROCON 2016, Dec 8–10, Bangalore, PPPV 79, p 162 Waterworth HE (1971) Physical properties and host ranges of viruses latent in and mechanically transmitted from jasmine. Phytopathology 61:228–230 Waterworth HE (1975) Purification of Arabis mosaic virus isolated from a jasmine plant introduction. Phytopathology 65:927–928 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98: 352–354

Jatropha spp. Family: Euphorbiaceae

Weed host

African cassava mosaic virus Taxonomic position Genus: Begomovirus

(ACMV)

Family: Geminiviridae

Jatropha spp.

1295

ACMV infection in plants of Jatropha curcas was reported from Kenya (Ramkat et al. 2011). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ACMV, refer to Manihot esculenta.

(CroYVMV)

Croton yellow vein mosaic virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

CroYVMV infection in plants of Jatropha gossypifolia was reported from India (Snehi et al. 2011b). The virus-infected jatropha plants exhibit symptoms of yellow vein mosaic, leaf deformation, vein swelling, and stunting. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of CroYVMV, refer to Croton bonplandianum (Syn.) Croton sparsiflorus.

(CMV)

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

J

Family: Bromoviridae

CMV infection in plants of Jatropha curcas was reported from India (Raj et al. 2008a, b). The virus-infected Jatropha plants exhibit severe mosaic disease accompanied by yellow spotting. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

East African cassava mosaic virus Taxonomic position Genus: Begomovirus

(EACMV)

Family: Geminiviridae

EACMV infection in plants of Jatropha curcas was reported from Kenya (Ramkat et al. 2011). The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of EACMV, refer to Manihot esculenta.

Indian cassava mosaic virus Taxonomic position Genus: Begomovirus

(ICMV)

Family: Geminiviridae

ICMV infection in plants of Jatropha curcas was reported from India (Gao et al. 2010; Snehi et al. 2012). The virus-infected jatropha plants exhibit mosaic symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of ICMV, refer to Manihot esculenta.

1296

Jatropha spp.

(JLCGV)

Jatropha leaf curl Gujarat virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution JLCGV infection in plants of Jatropha curcas was reported from India (Agarwal et al. unpublished – KM411359). Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2758 nt (KM411359) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

(JLCuV)

Jatropha leaf curl virus Synonyms Jatropha mosaic Lucknow virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution JLCuV infection in plants of Jatropha spp. was reported from India (Srivastava et al. 2015a). Symptoms and host(s) The virus-infected jatropha plants exhibit foliar yellow mosaic symptoms. Transmission JLCuV was shown to be transmitted by the whitefly Bemisia tabaci, the vector of begomoviruses. It is likely that, in common with other begomoviruses, JLCuV will be transmitted in a circulative, non-propagative manner. Similarly it is likely that the virus will prove not to be either mechanically or seed-transmitted. Experimentally the JLCuV was also whitefly transmitted to Nicotiana glutinosa and Solanum lycopersicum plants inducing yellow mosaic symptoms. The virus was also introduced into J. integerrima, J. podagrica, J. multifida, S. lycopersicum, N. glutinosa, and N. benthamiana plants by Agrobacterium-mediated inoculation of the cloned virus inducing yellow mosaic symptoms (Srivastava et al. 2015a). Virion properties and genome The structure of the virions of JLCuV has not been investigated. In common with all geminiviruses, the virions of JLCuV are likely geminate (twinned quasi-icosahedra). JLCuV is a typical old-world monopartite begomovirus. The genome of JLCuV consists of a single circular molecule of single-stranded DNA of 2743 nt (EU798996 = NC_011268; GU451249,

Jatropha spp.

1297

KC513823, KF652077, KF652078) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes of JLCuV encode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated.

Jatropha leaf yellow mosaic virus Taxonomic position Genus: Begomovirus

(JLYMV)

Family: Geminiviridae

Geographical distribution JLYMV infection in plants of Jatropha curcas was reported from India (Srivastava et al. 2015b). Symptoms and host(s) The virus-infected jatropha plants exhibit leaf yellow mosaic symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2744 nt (JN698954) (Briddon 2001; Brown et al. 2015; Srivastava et al. 2015b; Zerbini et al. 2017).

Jatropha mosaic India virus Taxonomic position Genus: Begomovirus

(JMINV)

Family: Geminiviridae

Geographical distribution JMINV infection in plants of Jatropha podagrica was reported from India (Narayana et al. 2007; Snehi et al. 2013, 2016; Srivastava et al. 2015b). Symptoms and host(s) The virus-infected jatropha plants exhibit severe foliar yellow mosaic symptoms. Transmission The transmission of JMINV has not been investigated. It is likely that, in common with other begomoviruses, JMINV is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of JMINV has not been investigated. In common with all geminiviruses, the virions of JMINV are likely geminate (twinned quasi-icosahedra). JMINV is a typical old-world monopartite begomovirus; no DNA-B or betasatellite was associated with the isolate of the virus identified so far. The genome of JMINV consists of a single circular

J

1298

Jatropha spp.

molecule of single-stranded DNA of 2740 nt (HM230683 = NC_038449; JN135236, JN698954) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes of JMINV isolates encode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated.

(JMNV)

Jatropha mosaic Nigeria virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution JMNV infection in plants of Jatropha curcas was reported from Nigeria (Kashina et al. 2013). Symptoms and host(s) The virus-infected jatropha plants exhibit symptoms of severe mosaic, mottling, and blistering of leaves. Transmission The transmission of JMNV has not been investigated. It is likely that, in common with other begomoviruses, JMNV is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of JMNV has not been investigated. In common with all geminiviruses, the virions of JMNV are likely geminate (twinned quasi-icosahedra). JMNV is a typical old-world monopartite begomovirus; no DNA-B or betasatellite was associated with the isolate of the virus identified so far. The genome of JMNV consists of a single circular molecule of single-stranded DNA of 2781 nt (JX025358 = NC_019034; JX025359, JX025360) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes of JMNV isolates encode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated.

Jatropha mosaic virus Taxonomic position Genus: Begomovirus

(JMV)

Family: Geminiviridae

Geographical Distribution JMV infection was first reported in plants of Jatropha gossypifolia from Puerto Rico by Bird (1957). The virus infection in Jatropha spp. was reported from Karnataka (India), Kenya, Florida, Jamaica, Dominican Republic, and Puerto Rico (Kim et al. 1986; Jayanna 2006; Narayana et al. 2006a; Roye et al. 2006; Polston et al. 2014; Simmonds-Gordon et al. 2014; Melgarejo et al. 2015).

Jatropha spp.

1299

Symptoms and host(s) The virus-infected jatropha plants exhibit foliar yellow mosaic symptoms; virus-infected Jatropha curcas plants exhibit symptoms of mosaic, reduced leaf size, leaf distortion, blistering, and stunting of the plants. This virus also infects solanaceous hosts like tobacco and Nicotiana benthamiana (Melgarejo et al. 2015). Transmission The transmission of JMV has not been investigated. It is likely that, in common with other begomoviruses, JMV is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner (Narayana et al. 2006b). The virus is mechanically transmissible to some hosts. Isolates of JMV from the Dominican Republic were mechanically transmissible to Nicotiana benthamiana and common bean (Phaseolus vulgaris) leading to stunting, severe foliar crumpling, deformation, and yellow mosaic symptoms. Additionally, isolates from the Dominican Republic were infectious to N. benthamiana (leading to stunting, severe foliar crumpling, deformation, and yellow mosaic symptoms), N. glutinosa (symptomless), N. tabacum (mosaic, leaf curling, leaf deformation, and stunting dependent upon the variety), and a Jatropha spp. (mosaic, leaf curling, leaf deformation, and stunting) by Agrobacterium-mediated inoculation of cloned viral components (Melgarejo et al. 2015). Virion properties and genome The structure of the virions of JMV has not been investigated. In common with all geminiviruses, the virions of JMV are likely geminate (twinned quasi-icosahedra). JMV is typical of bipartite begomoviruses native to the New World. The genome consists of two circular, single-stranded DNA components. DNA-A consists of 2610 nt (KF723258 = NC_024012; KF723259, KF723260) and DNA-B of 2588 nt (KF723261 = NC_024013; KF723262) (Briddon 2001; Polston et al. 2014; Simmonds-Gordon et al. 2014; Brown et al. 2015; Zerbini et al. 2017). Typical of the majority of bipartite begomoviruses native to the New World, the DNA-A component of JMV encodes five genes, one in the virion-sense and four in the complementary-sense, whereas the DNA-B encodes one gene in each orientation. The expression and function of these genes have not been investigated for JMV.

Jatropha yellow mosaic virus Taxonomic position Genus: Begomovirus

(JYMV)

Family: Geminiviridae

Geographical distribution JYMV infection in plants of Jatropha gossypifolia was reported from India (Snehi et al. 2011a). Symptoms and host(s) The virus-infected jatropha plants exhibit yellow mosaic symptoms and reduction in leaf size. Transmission The transmission of JYMV has not been investigated. It is likely that, in common with other begomoviruses, JYMV is transmitted by the whitefly Bemisia tabaci in a circulative, non-propagative manner.

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Jatropha spp.

Virion properties and genome The structure of the virions of JYMV has not been investigated. In common with all geminiviruses the virions of JYMV will likely be geminate (twinned quasi-icosahedra). JYMV is a typical Old World monopartite begomovirus; no DNA B or betasatellite was associated with the one isolate of the virus identified so far. The genome of JYMV consists of a single circular molecule of single-stranded DNA (DNA-A) of 2757 nt (FJ177030 = NC_011309) (Briddon 2001; Snehi et al., 2011a; Brown et al. 2015; Zerbini et al. 2017). The characterised genome of JYMV encodes the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes has not been investigated.

(PaLCuV)

Papaya leaf curl virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

The virus infection in plants of Jatropha integerrima was reported from India (Snehi et al. 2013, 2016). The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The virus is not mechanically sap-transmissible. For more details of PaLCuV, refer to Carica papaya.

Tomato leaf curl Patna virus Taxonomic position Genus: Begomovirus

(ToLCPatV)

Family: Geminiviridae

ToLCPatV infection in plants of Jatropha multifida was reported from India (Snehi et al. 2013, 2016). The virus-infected jatropha plants exhibit leaf curling symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. For more details of ToLCPatV, refer to Solanum lycopersicum.

References Bird J (1957) A whitefly-transmitted mosaic of Jatropha gossypiifolia. Technical Paper of the Agricultural Experiment Station of Puerto Rico, 22:1–35 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer Index of Viruses. Springer New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Gao SQ, Qu J, Chua N-H, Ye J (2010) A new strain of Indian cassava mosaic virus causes a mosaic disease in the biodiesel crop Jatropha curcas. Arch Virol 155:607–612 Jayanna K (2006) Studies on Jatropha mosaic virus disease. MSc thesis submitted to the Department of Plant Pathology, College of Agriculture, Dharwad University of Agricultural Sciences, Dharwad-580005, pp 80 Kashina BD, Alegbejo MD, Banwo OO, Nielsen SL, Nicolaisen M (2013) Molecular identification of a new begomovirus associated with mosaic disease of Jatropha curcas L. in Nigeria. Arch Virol 158:511–514 Kim KS, Bird J, Rodriguez RL, Martin EM, Escudero J (1986) Ultrastructural studies of Jatropha gossypifolia infected with jatropha mosaic virus, a whitefly-transmitted geminivirus. Phytopathology 76:80–85 Melgarejo TA, Kon T, Gilbertson RL (2015) Molecular and biological characterization of distinct strains of Jatropha mosaic virus from the Dominican Republic reveal a potential to infect crop plants. Phytopathology 105:141–153

Juglans regia (Walnut)

1301

Narayana DSA, Shankarappa KS, Govindappa MR, Prameela HA, Gururaj Rao MR, Rangaswamy KT (2006a) Natural occurrence of Jatropha mosaic virus disease in India. Curr Sci 91:585–586 Narayana DSA, Shankarappa KS, Prameela HA, Keshavamurthy KV, Rangaswamy KT (2006b) PCR detection of Jatropha mosaic virus in whitefly Bemisia tabaci and Dodder Cuscuta subilclusa. Indian J Virol 17:144–145 Narayana DSA, Rangaswamy KT, Shankarappa KS, Maruthi MN, Lakshminarayana Reddy CN, Rekha AR, Keshava Murthy KV (2007) Distinct begomoviruses closely related to cassava mosaic viruses cause Indian jatropha mosaic disease. Int J Virol 3:1–11 Polston JE, Londono MA, Capobianco H (2014) The complete genome sequence of New World Jatropha mosaic virus. Arch Virol 159(11):3131–3136 Raj SK, Kumar S, Snehi SK, Pathre U (2008a) First report of Cucumber mosaic virus on Jatropha curcas in India. Plant Dis 92:171 Raj SK, Snehi SK, Kumar S, Khan MS, Pathre U (2008b) First molecular identification of a Begomovirus in India that is closely related to Cassava mosaic virus and causes mosaic and stunting of Jatropha curcas L. Aust Plant Dis Notes 3:69–72 Ramkat RC, Alberto C, Fatemeh M, Margit L (2011) Occurrence of African cassava mosaic virus (ACMV) and East African cassava mosaic virus–Uganda (EACMV-UG) in Jatropha curcas. BMC Proc 5:P93 Roye M, Collins S, Maxwell DP (2006) The first report of a begomovirus associated with the common weed Jatropha gossypifolia in Jamaica. Plant Pathol 55:286 Simmonds-Gordon RN, Collins-Fairclough AM, Stewart CS, Roye ME (2014) First report of a complete genome sequence for a begomovirus infecting Jatropha gossypifolia in the Americas. Arch Virol 159:2815–2818 Snehi SK, Khan MS, Raj SK, Prasad V (2011a) Complete nucleotide sequence of Croton yellow vein mosaic virus and DNA- b associated with yellow vein mosaic disease of Jatropha gossypifolia in India. Virus Genes 43:93–101 Snehi SK, Raj SK, Khan MS, Prasad V (2011b) Molecular identification of a new begomovirus associated with yellow mosaic disease of Jatropha gossypifolia in India. Arch Virol 156:2303–2307 Snehi SK, Srivastava A, Raj SK (2012) Biological characterization and complete genome sequence of a possible strain of Indian cassava mosaic virus associated with mosaic disease of Jatropha curcas in India. J Phytopathol 160:547–553 Snehi SK, Srivastava A, Kumar S, Raj SK (2013) Molecular detection and identification of Begomoviruses affecting important ornamental plants in India. Indian J Virol 24:142 Snehi SK, Raj SK, Prasad V, Singh V (2016) Molecular detection and identification of Begomovirus isolates associated with mosaic disease of ornamental Jatropha species from India. Int J Bact Virol 1:004 Srivastava A, Kumar S, Jaidi M, Raj SK (2015a) Characterization of a novel begomovirus associated with yellow mosaic disease of three ornamental species of Jatropha grown in India. Virus Res 201:41–49 Srivastava A, Kumar S, Jaidi M, Raj SK (2015b) Molecular characterization of a new begomovirus associated with leaf yellow mosaic disease of Jatropha curcas in India. Arch Virol 160:1359–1362 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Geminiviridae. J Gen Virol 98:131–133

Juglans regia (Walnut) Family: Juglandaceae

Cherry leaf roll virus Taxonomic position Genus: Nepovirus

Trees/Shrubs

(CLRV)

Family: Secoviridae

CLRV infection in trees of Juglans regia was reported from the UK, Bulgaria, England, France, Hungary, Romania, Spain, the USA (California and Oregon), Italy, Turkey, Syria, and New Zealand (Quacquarelli and Savino 1977; Savino et al. 1977; Cooper 1980; Cooper and Edwards 1980; Mircetich et al. 1980; Mircetich and Rowhani 1984; Rowhani et al. 1985; Baumgartenerova and Slovaka 1995; Borja et al. 1995; Lazarova-Topchiiska 1995; Ozturk et al. 2008; Al-Chaabi and Ismaeil 2009; von Bargen et al. 2009; Ferretti et al. 2017). The virus-infected walnut trees exhibit chlorotic mosaic, spots,

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Justicia brandegeana (Shrimp plant)

rings, and yellow line pattern symptoms. The virus is transmitted by nematode vectors Xiphinema coxi, X. diversicaudatum, and X. vuittenezi in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants. The virus is seed transmitted in Juglans regia to the extent of 52.3% (Kolber et al. 1982; Mircetich et al. 1982). There is evidence of CLRV transmission through pollen. For more details of CLRV, refer to Prunus avium.

References Al-Chaabi S, Ismaeil F (2009) First report of the cherry strain of Cherry leaf roll virus on walnut and pecan trees in Syria. J Plant Pathol 91:502 von Bargen S, Grubits E, Jalkanen R, Büttner C (2009) Cherry leaf roll virus – an emerging virus in Finland? Silva Fenn 43:727–738 Baumgartenerova H, Slovaka L (1995) First report of Walnut ringspot caused by Cherry leaf roll virus from Slovakia. Z Pflanzenkrankh Pflanz 102:435–440 Borja M, Sanchez F, Rowhani A, Bruening G, Ponz F (1995) Long, nearly identical untranslated sequences at the 30 terminal regions of the genomic RNA’s of Cherry leafroll virus (Walnut strain). Virus Genes 10:245–252 Cooper JI (1980) The prevalence of Cherry leaf roll virus in Juglans regia in the United Kingdom. Acta Phytopathol Acad Sci Hung 15:139–145 Cooper JI, Edwards ML (1980) Cherry leaf roll virus in Juglans regia in the United Kingdom. Forestry 53:41–50 Ferretti L, Corsi B, Luongo L, Dal Cortivo C, Belisario A (2017) A survey of Cherry leaf roll virus in intensively managed grafted English (Persian) walnut trees in Italy. J Plant Pathol 99:423–427 Kolber M, Nemeth M, Szentivanyi P (1982) Routine testing of English walnut mother trees and group testing of seeds by ELISA for direction of Cherry leaf roll virus infection. Acta Hortic 130:161–169 Lazarova-Topchiiska M (1995) Identification of the cherry leaf roll virus in Persian walnut (Juglans regia L.) tissue. Plant Sci 32(7–8):99–108 Mircetich SM, Rowhani A (1984) The causal relationship of Cherry leafroll virus and blackline disease of English walnut trees. Phytopathology 74:423–428 Mircetich SM, Sanborn RR, Ramos DE (1980) Natural spread, graft-transmission and possible etiology of walnut blackline disease. Phytopathology 70:962–968 Mircetich S, Rowhani A, Cucuzza J (1982) Seed and pollen transmission of Cherry leaf roll virus (CLRV-W), the causal agent of the black-line (BL) disease of walnut trees. Phytopathology 72:988 Ozturk MO, Sipahioglu HM, Ocak M, Usta M (2008) Cherry leaf roll virus in Juglans regia in the Lake Van basin of Turkey. J Plant Pathol 90:75–79 Quacquarelli A, Savino V (1977) Cherry leaf roll virus in walnut (Juglans regia): II. Distribution in Apulia and transmission through seed. Phytopathol Mediterr 16:154–156 Rowhani A, Mircetich SM, Shepherd RJ, Cucuzza JD (1985) Serological detection of Cherry leaf roll virus in English walnut trees. Phytopathology 75:48–52 Savino V, Quacquarelli A, Gallitelli A, Gallitelli D, Piazzolla P, Martelli G (1977) Cherry leaf roll virus in walnut. I. Identification and characterization. Phytopathol Mediterr 16:96–102

Justicia brandegeana (Shrimp plant) Family: Acanthaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

Justicia brandegeana (Shrimp plant)

1303

CMV infection in plants of Justicia brandegeana was reported from Iraq (Adhab and Al-Ani 2011). The virus-infected shrimp plants exhibit symptoms of chlorotic lesions on the leaves. The virus is transmitted by aphid vectors, Myzus persicae and Aphis gossypii, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Justicia brandegeana was reported from Greece (Chatzivassiliou et al. 2000). The virus-infected shrimp plants exhibit symptoms of chlorotic and necrotic foliar rings and floral malformation and necrosis. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Adhab MA, Al-Ani RA (2011) Amaryllis and shrimp plant are secondary hosts of Cucumber mosaic cucumovirus (CMV) in Iraq. Agric Biol J N Am 2:872–875 Chatzivassiliou EK, Livieratos I, Jenser G, Katis NI (2000) Ornamental plants and thrips populations associated with Tomato spotted wilt virus in Greece. Phytoparasitica 28:257–264

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Kalanchoe blossfeldiana (Kalanchoe) Family: Crassulaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV) Family: Bromoviridae

CMV infection in plants of Kalanchoe blossfeldiana was reported from Iran (Shahmohammadi et al. 2015) although no specific symptoms were described. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV occurs wherever Kalanchoe blossfeldiana is grown (Mertelik et al. 2002). The virus-infected kalanchoe plants show yellowish chlorotic rings, spots, or patterns on leaves; some leaves may wilt. The virus is transmitted by the thrips vector, Frankliniella occidentalis, in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Kalanchoe latent virus Taxonomic position Genus: Carlavirus

(KLV) Family: Betaflexiviridae

© Springer Nature India Private Limited 2019 K. S. Sastry et al., Encyclopedia of Plant Viruses and Viroids, https://doi.org/10.1007/978-81-322-3912-3

1306

Kalanchoe blossfeldiana (Kalanchoe)

Geographical distribution This virus was first reported in plants of Kalanchoe blossfeldiana from the USA (Hearon 1981, 1982). KLV occurs wherever K. blossfeldiana is grown. Symptoms and host(s) Hearon (1984) has isolated KV-1 and KV-2 strains of KLV from the USA, and this virus is known to cause symptomless infection in several cultivars of K. blossfeldiana. KLV has also been reported to infect Graptopetalum paraguayense in Italy (Sorrentino et al. 2017). Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus spreads through infected cuttings used as planting materials. Virus is transmissible by mechanical sap-inoculation to a few members of the family Chenopodiaceae (Hearon 1982). Virion properties and genome The virions are flexuous filaments about 620–650 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8517 nt (FJ531634 = NC_013006) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Nicolaisen and Nielsen 2001; Adams et al. 2004; Dinesen et al. 2009).

Kalanchoe mosaic virus Taxonomic position Genus: Potyvirus

(KMV)

Family: Potyviridae

Geographical distribution KMV infection in plants of Kalanchoe blossfeldiana was reported from the USA and European countries (Husted and Bech 1996; Bjorke et al. 2012). Symptoms and host(s) This virus typically produces uneven green color in the leaves often described as “green islands,” curling of the leaves, and reduced growth (Husted and Bech 1996). KMV has also been reported to infect Graptopetalum paraguayense in Italy (Sorrentino et al. 2017). Transmission The virus is transmitted by an aphid vector Myzus persicae in a non-persistent manner. It is also transmissible by mechanical sap-inoculation to a very limited number of families. The use of virusinfected vegetative propagative material is the primary mode of virus spread. The virus is also grafttransmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, with a clear modal length of 724 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA (Husted et al. 1994; Revers and Garcia 2015). A partial genome sequence of 9462 nt (KY385304) lacking part of the 50 region is available (Wylie et al. 2017).

Kalanchoe blossfeldiana (Kalanchoe)

1307

Kalanchoe top-spotting virus Taxonomic position Genus: Badnavirus

(KTSV)

Family: Caulimoviridae

Geographical distribution KTSV infection in plants of Kalanchoe blossfeldiana was reported from Europe and North America (Hearon and Locke 1984; Yang et al. 2005). Symptoms and host(s) Infection is characterized by the appearance of numerous yellow spots on the leaves of virus-infected kalanchoe plants. Spots usually fade as the leaves expand, but circular depressions often develop on the upper leaves surface at sites that formerly showed chlorotic spots. Transmission The virus is transmissible by mechanical inoculation to a limited number of hosts and by the citrus mealybug, Planococcus citri, in a semi-persistent manner. The virus is retained after molts, but does not multiply in the mealybug and is not transmitted through progeny (Hearon and Locke 1984). The natural spread of this virus in commercially grown kalanchoe, occurs primarily by vegetative propagation of infected stock plants. The virus is seed-transmitted to the extent of 60–90% and is also transmitted in pollen. Virion properties and genome The virions are bacilliform in shape with a diameter of 30 nm, and modal particle length is 130 nm. The genome is a monopartite, circular, double-stranded DNA of 7591 bp with a single-strand discontinuity at one site in each strand (AY180137 = NC_004540) (Lockhart and Ferji 1988; Yang 1995; Yang et al. 2005; Olszewski and Lockhart 2011; Bhat et al. 2016).

Sonchus yellow net nucleorhabdovirus Taxonomic position Genus: Nucleorhabdovirus

(SYNV)

Family: Rhabdoviridae

SYNV infection in plants of Kalanchoe blossfeldiana was reported from the Netherlands (Bouwen et al. 2002). The virus-infected kalanchoe plants exhibit chlorotic spot symptoms. The virus is transmitted by aphid vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of SYNV, refer to Lactuca sativa.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV is worldwide in distribution (Verhoeven 1992; German et al. 1992; Hausbeck et al. 1992; Uga and Tsuda 2005). Virus-infected kalanchoe plants show diffuse chlorotic spots on young leaves. Sometimes chlorosis and necrosis are seen on leaves, petioles, and stems of many cultivars; the

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Kalanchoe blossfeldiana (Kalanchoe)

environmental factors have an influence on symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a number of test plants. Planting of infected material is responsible for maximum spread of this virus disease. For more details of TSWV, refer to Solanum lycopersicum.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Bhat AI, Hohn T, Selvarajan R (2016) Badnaviruses: the current global scenario. Viruses 8(6):177 Bjorke L, Willumsen K, Skjeseth G, Haugslien S, Blystad D-R (2012) The effect of Kalanchoe mosaic virus on growth and development on four cultivars of Kalanchoe blossfeldiana. Bioforsk Fokus 7(9):9 Bouwen I, Schoen CD, van Balen E, van der Vlugt RAA (2002) Kalanchoe blossfeldiana, a new host for Sonchus yellow net virus. Acta Hortic 568:59–60 Dinesen M, Lundmark M, Albrechtsen M (2009) Complete genome sequences of two isolates of Kalanchoe latent virus. Arch Virol 154:1173–1175 German TL, Ullman DE, Moyer JW (1992) Tospoviruses: diagnosis, molecular biology, phylogeny, and vector relationships. Annu Rev Phytopathol 30:315–348 Hausbeck MK, Welliver RA, Derr MA, Gildow FE (1992) Tomato spotted wilt virus survey among green house ornamentals in Pennsylvania. Plant Dis 76:795–800 Hearon SS (1981) Detection of viruses in ultrastructural studies of naturally infected Kalanchoe blossfeldiana. Phytopathology 71:767 Hearon SS (1982) A carlavirus from Kalanchoe blossfeldiana. Phytopathology 72:838–844 Hearon SS (1984) Comparison of two strains of Kalanchoe latent virus, carlavirus group. Phytopathology 74:670–676 Hearon SS, Locke JC (1984) Graft, pollen, and seed transmission of an agent associated with top spotting in Kalanchoe blossfeldiana. Plant Dis 68:347–350 Husted K, Bech K (1996) Stability, host range, and distribution of Kalanchoe mosaic potyvirus in Kalanchoe blossfeldiana. Plant Dis 80:211–214 Husted K, Bech K, Albrechtsen M, Borkhardt B (1994) Identification, partial sequencing and detection of a potyvirus from Kalanchoe blossfeldiana. Phytopathology 84:161–164 Lockhart BEL, Ferji Z (1988) Purification and mechanical transmission of Kalanchoe top spotting-associated virus. Acta Hortic 234:72–78 Mertelik J, Mokra V, Gotzova B, Gabrielova S (2002) Occurrence and identification of impatiens necrotic spot tospovirus in the Czech Republic. Acta Hortic 568:79–83 Nicolaisen M, Nielsen SL (2001) Analysis of the triple gene block and coat protein sequences of two strains of Kalanchoe latent carlavirus. Virus Genes 22(3):265–270 Olszewski NE, Lockhart B (2011) Badnavirus. Caulimoviridae. In: The Springer Index of Viruses. Springer New York, pp 263–269. https://doi.org/10.1007/978-0-387-95919-1_40 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Sorrentino R, Marais A, Faure C, Thell S, Alioto D, Candresse T (2017) First report of Kalanchoe mosaic virus and Kalanchoe latent virus infecting ghost plant (Graptopetalum paraguayense) in Italy. Plant Dis 101:1560 Shahmohammadi N, Dizadji A, Habibi MK, Nateqi M (2015) First report of Cucumber mosaic virus infecting Bougainvillea spectabilis, Coleus blumei, Kalanchoe blossfeldiana and Zinnia elegans in Iran. J Plant Pathol 97(2):394 Uga H, Tsuda S (2005) A one-step reverse transcription-polymerase chain reaction system for the simultaneous detection and identification of multiple tospovirus infections. Phytopathology 95:166–171 Verhoeven JTJ (1992) Tomato spotted wilt virus: ecological aspects in ornamental crops in the Netherlands from 1989 up to 1991. Acta Hortic 377:175–182 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, and ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354 Yang Z (1995) An analysis of the nucleotide sequence and genomic organization of Kalanchoe top spotting virus. MS thesis, University of Minnesota, St. Paul Yang Z, Nicolaisen M, Olszewski NE, Lockhart BEL (2005) Sequencing, improved detection, and a novel form of Kalanchoe top-spotting virus. Plant Dis 89:298–302

Kennedia spp. (Kennedia rubicunda, K. prostrata)

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Kalimeris indica (Indian aster) Family: Asteraceae

Medicinal

Papaya leaf curl virus

(PaLCuV)

Taxonomic position Genus: Begomovirus

Family: Geminiviridae

PaLCuV infection in plants of Kalimeris indica was reported from India (Srivastava et al. 2013). The virus-infected Indian aster plants exhibit yellow vein symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not mechanically saptransmissible. For more details of PaLCuV, refer to Carica papaya.

Potato virus Y

(PVY)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

PVY infection in plants of Kalimeris indica was reported from China (Wang et al. 2012). The virusinfected Indian aster plants exhibit symptoms of diffuse chlorotic and necrotic spots on the leaves. The virus is transmitted by aphid vectors in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation to a large number of hosts and also by grafting. For more details of PVY, refer to Solanum tuberosum.

References Srivastava A, Raj SK, Kumar S, Snehi SK (2013) New record of Papaya leaf curl virus and Ageratum leaf curl beta satellite associated with yellow vein disease of aster in India. New Dis Rep 28:6 Wang F, Wu YH, Gao ZL, Zhou BG, Guo L (2012) First report of Potato virus Y in Kalimeris indica in China. Plant Dis 96:1827

Kennedia spp. (Kennedia rubicunda, K. prostrata) Family: Fabaceae

Ornamental

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

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1310

Kennedia spp. (Kennedia rubicunda, K. prostrata)

BYMV infection in plants of Kennedia prostrata was reported from Australia (McKirdy et al. 1994; Webster et al. 2007). The virus-infected kennedia plants exhibit mosaic and stunting symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Kennedya yellow mosaic virus Taxonomic position Genus: Tymovirus

(KYMV)

Family: Tymoviridae

Geographical distribution KYMV was first reported in plants of Kennedya rubicunda in Australia and Tasmania (Dale et al. 1975; Dale and Gibbs 1976). Symptoms and host(s) The virus-infected kennedia plants exhibit bright blotchy yellow mosaic symptoms. Transmission The virus is transmitted by beetle vectors in a semi-persistent manner. The virus is mechanically saptransmissible to a few members of Papilionaceae and Solanaceae (Dale and Gibbs 1976). The virus is not transmissible by contact between plants or through pollen and seed. Virion properties and genome The virions are non-enveloped, isometric capsid with T = 3 icosahedral symmetry, about 30 nm in diameter. The genome consists of a single molecule of positive-sense, ssRNA of 6362 nt (D00637 = NC_001746). The 30 -terminus has a tRNA-like structure (Ding et al. 1990; Martelli et al. 2002).

References Dale J, Gibbs AJ (1976) Kennedya yellow mosaic virus: another tymovirus. Aust J Biol Sci 29:397–403 Dale J, Gardiner JE, Gibbs A (1975) Viruses in Kennedya rubicunda. Aust Plant Pathol Soc Newsl 4:13–14 Ding S, Keese P, Gibbs A (1990) The nucleotide sequence of the genomic RNA of Kennedya yellow mosaic tymovirusJervis Bay isolate: relationships with potex- and carlaviruses. J Gen Virol 71(PT 4):925–931 Martelli GP, Sabanadzovic S, Abou-Ghanem Sabanadzovic N, Edwards MC, Dreher T (2002) The family Tymoviridae. Arch Virol 147(9):1837–1846 McKirdy SJ, Coutts BA, Jones RAC (1994) Occurrence of Bean yellow mosaic virus in subterranean clover pastures and perennial native legumes. Aust J Agr Res 45:183–194 Webster CG, Coutts BA, Jones RAC, Jones MGK, Wylie SJ (2007) Virus impact at the interface of an ancient ecosystem and a recent agroecosystem: studies on three legume-infecting potyviruses in the southwest Australian floristic region. Plant Pathol 56:729–742

Kohleria spp.

1311

Kohleria spp. Family: Gesneriaceae

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

Ornamental

(TMV) Family: Virgaviridae

TMV was reported from cultivated Kohleria spp. in the USA, but not from any wild-collected plants (Zettler and Nagel 1983). The symptoms on the foliage of infected Kohleria plants were inconspicuous. There is no known vector for this virus. The virus is transmissible by mechanical sap-inoculation, and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

References Zettler FW, Nagel J (1983) Infection of cultivated Gesneriads by two strains of Tobacco mosaic virus. Plant Dis 67:1123–1125

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Laburnum spp. Family: Fabaceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

Family: Secoviridae

ArMV infection in plants of Laburnum spp. was reported from the UK and Germany (Schmelzer 1962/ 1963; Cooper and Sweet 1976). The virus-infected laburnum plants exhibit symptoms of stunted and chlorotic shoots. This virus is transmitted by nematode vectors (Xiphinema spp.) in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV was isolated from plants of Laburnum alpinum in the USA showing conspicuous vein-banding but was not demonstrated to be the cause of the symptoms observed (Brierley and Smith 1954). The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

References Brierley P, Smith FF (1954) New records of virus diseases of ornamental plants. Plant Dis Reptr 38:739–741 Cooper JI, Sweet JB (1976) The detection of viruses with nematode vectors in six woody hosts. Forestry 49:73–78 Schmelzer K (1962/63) Untersuchungen an viren der zier und wildgeholze 2. Mitteilung: Virosen an Forsythia, Lonicera, Ligustrum, und Laburnum. Phytopathol Z 46:105–138 © Springer Nature India Private Limited 2019 K. S. Sastry et al., Encyclopedia of Plant Viruses and Viroids, https://doi.org/10.1007/978-81-322-3912-3

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Lachenalia spp.

Lachenalia spp. Family: Asparagaceae

Ornamental

Freesia sneak ophiovirus Taxonomic position Genus: Ophiovirus

(FreSV)

Family: Aspiviridae

FreSV infection in plants of Lachenalia cultivars was reported from South Africa (Vaira et al. 2007). The virus-infected lachenalia plants exhibited fine chlorotic streaking and occasional gray flecking. The virus is transmitted by the soil-borne fungus Olpidium brassicae and also by mechanical sap-inoculation. For more details of FreSV, refer to Freesia spp.

Ornithogalum mosaic virus Taxonomic position Genus: Potyvirus

(OrMV)

Family: Potyviridae

OrMV-infected plants of Lachenalia spp. were first reported from South Africa (Burger and von Wechmar 1988). The virus-infected lachenalia plants exhibited prominent chlorotic streaking, necrosis, and/or leaf deformation symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of OrMV, refer to Ornithogalum spp.

References Burger JT, von Wechmar MB (1988) Rapid diagnosis of Ornithogalum and Lachenalia viruses in propagation stock. Acta Hortic 234:31–38 Vaira AM, Kleynhans R, Hammond J (2007) First report of Freesia sneak virus infecting Lachenalia cultivars in South Africa. Plant Dis 91:770

Lactuca indica (Indian lettuce) Family: Asteraceae

Broad bean wilt virus 2 Taxonomic position Genus: Fabavirus

Weed host

(BBWV-2)

Family: Secoviridae

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BBWV-2 infection in plants of Lactuca indica was reported from China (Wu et al. 2010). The virus-infected Indian lettuce plants exhibit mottled leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BBWV-2, refer to Vicia faba.

References Wu XY, Wu JX, Cheng XF (2010) First report of natural infection of Broad bean wilt virus 2 on Lactuca Indica. J Plant Pathol 92:S4.119

Lactuca sativa (Lettuce) Family: Asteraceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Vegetable

(AMV)

Family: Bromoviridae

AMV occurs wherever plants of Lactuca sativa are grown (Stone and Nelson 1966; Fletcher 2001; Moreno et al. 2004; Blancard et al. 2006). The virus-infected lettuce plants produce symptoms which including bright yellow mosaic or calico patterns on leaves. Older leaves develop a yellow mottle with enlarged veins. Infected plants may be stunted. The virus is transmitted by aphid vectors in a nonpersistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of AMV, refer to Medicago sativa.

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

ArMV infection in plants of Lactuca sativa was reported from New Zealand (Mossop et al. 1983). The virus-infected lettuce plants either remained symptomless or showed mild chlorosis. The virus is transmitted by a nematode vector in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Beet mosaic virus Taxonomic position Genus: Potyvirus

(BtMV)

Family: Potyviridae

L

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Lactuca sativa (Lettuce)

BtMV was reported to infect plants of Lactuca sativa in China, during 2004–2006 (Wang et al. 2008). The affected lettuce plants showed mosaic, puckering and severe stunting symptoms. The virus is transmitted by aphid vectors in a non-persistent manner; by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants. For more details of BtMV, refer to Beta vulgaris.

Beet pseudoyellows virus Taxonomic position Genus: Crinivirus

(BPYV)

Family: Closteroviridae

BPYV was reported to infect plants of Lactuca sativa in England (Coffin and Coutts 1990). The virusinfected lettuce plants were stunted; leaves developed severe interveinal chlorosis and slight downward curling. The virus is transmitted by the greenhouse whitefly (Trialeurodes vaporariorum) in a semipersistent manner. The virus is not transmissible by mechanical sap-inoculation, and also not by contact between plants. For more details of BPYV, refer to Beta vulgaris.

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

BWYVoccurs wherever Lactuca sativa is grown (Zink and Duffus 1972; Tomlinson and Walker 1973; Marco 1984; Lot and Maury-Chovelon 1985; Timmerman et al. 1985; Moreno et al. 2004; Xiang et al. 2008). On lettuce, this virus rarely produces symptoms until plants reach the rosette stage. With the advancement of the disease, the yellowing continues until all the oldest, lower leaves are bright yellow to sometimes almost white in color, with the main leaf veins remaining green. Yellowed leaves often have a thick, brittle texture. Yellowing can progress until the wrapper leaves adjacent to the head also turn yellow, and head color may be unacceptably light green. In most lettuce varieties, significant stunting or reduction in plant size does not occur. The virus is transmitted by a number of aphid species in a circulative, non-propagative manner. The virus is not transmitted through mechanical sap-inoculation, through seed, or through pollen. For more details of BWYV, refer to Beta vulgaris.

Beet yellow stunt virus Taxonomic position Genus: Closterovirus

(BYSV)

Family: Closteroviridae

BYSV infection in plants of Lactuca sativa was first reported from California (USA) and England (Duffus 1972). BYSV-infected lettuce plants exhibit symptoms of pronounced yellowing of older leaves; later, infected plants collapse prematurely and die. Internally, the phloem tissue of the stem and crown is brown and necrotic, which extends into the crown tissue. The virus is transmitted by aphids; the sow thistle aphid, Hyperomyces lactucae, is the most efficient vector, and the virus is transmitted in a semi-persistent manner. The virus is mechanically transmitted to number of host plants with difficulty,

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producing yellowing or reddening, and necrotic spots. The virus is not seed, pollen, and contact transmitted. For more details of BYSV, refer to Beta vulgaris.

Bidens mosaic virus Taxonomic position Genus: Potyvirus

(BiMV)

Family: Potyviridae

BiMV infection in plants of Lactuca sativa was reported from Brazil (Pavan et al. 2008; Suzuki et al. 2009). The virus-infected lettuce plants exhibit mosaic and foliar deformation symptoms. The virus is transmitted by aphid vectors, Myzus persicae, Aphis coreopsidis, and Dactynotus spp., in a nonpersistent manner, and also by mechanical sap-inoculation. For more details of BiMV, refer to Bidens spp.

Bidens mottle virus Taxonomic position Genus: Potyvirus

(BiMoV)

Family: Potyviridae

BiMoV is known to infect plants of Lactuca sativa in the USA and Taiwan (Purcifull et al. 1971; Chen and Lee 2012; Chen et al. 2012). The virus-infected lettuce plants exhibit symptoms of mosaic, leaf deformation and stunting of plants. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BiMoV, refer to Bidens spp.

Broad bean wilt virus 1 Taxonomic position Genus: Fabavirus

(BBWV-1)

Family: Secoviridae

BBWV-1 infects plants of Lactuca sativa in the USA and Spain (Bruckart and Lorbeer 1975; Belliure et al. 2009). The virus-infected lettuce plants exhibit symptoms of mottling of the young leaves and slight discoloration; normal plant growth is delayed. The virus is transmitted by aphid vectors, Myzus persicae and Aphis gossypii, in a non-persistent manner, and also by mechanical sap-inoculation (Belliure et al. 2009). For more details of BBWV-1, refer to Vicia faba.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMVoccurs in plants of Lactuca sativa in France, Brazil, Italy, Israel, Iran, Spain, Poland, Yemen, and Europe (Bruckart and Lorbeer 1975; Alhubaishi et al. 1987; Twardowicz-Jakusz and Zielinska 1997; Moreno et al. 2004; Pavan et al. 2008; Soleimani et al. 2011). The virus-infected lettuce plants exhibit symptoms of leaf mottling, severe roughness of the leaf, and occasional necrosis within the leaf tissue.

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Lactuca sativa (Lettuce)

Chlorotic mottling and crinkling of lamina can also be seen in infected plants. Plants are usually stunted if infected at an early stage of development. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(CABYV)

Family: Luteoviridae

CABYV infection in plants of Lactuca sativa was reported from Tunisia (Mnari-Hattab et al. 2009). The virus-infected lettuce plants show severe yellowing symptoms. The virus is transmitted by aphid vectors, Myzus persicae, M. euphorbiae, and Aphis gossypii, in a circulative, non-propagative manner, and is not transmissible by mechanical sap-inoculation. For more details of CABYV, refer to Cucumis melo.

Dandelion yellow mosaic virus Taxonomic position Genus: Sequivirus

(DaYMV)

Family: Secoviridae

DaYMV is known to infect plants of Lactuca sativa in the UK, the Netherlands, Poland, Chile, and Brazil (Kassanis 1947; Bos et al. 1983; Vetten et al. 1985; Twardowicz-Jakusz and Zielinska 1997; Krause-Sakate et al. 2005; Jadao et al. 2007). The young leaves of virus-infected lettuce plants become bronzed as a result of fine brown necrosis along the veins and interveinal areas. In the glasshouse, this is usually followed by chlorosis, dwarfing, and malformation of the whole plant. The virus is transmitted by aphid vectors Myzus ornatus, M. ascalonicus, and Aulacorthum solani in a semi-persistent manner, however, it is dependent on the presence of a helper virus in the genus Waikavirus and also by mechanical sap-inoculation. For more details of DaYMV, refer to Taraxacum officinale.

Groundnut ringspot orthotospovirus Taxonomic position Genus: Orthotospovirus

(GRSV)

Family: Tospoviridae

GRSV is known to infect plants of Lactuca sativa in Brazil (Chaves et al. 2001; Pavan et al. 2008). The virus-infected lettuce plants exhibit symptoms of circular necrotic spots and browning of the leaves. Eventually leaves becomes completely necrotic, leading to death of the plants. The virus is transmitted by thrips vectors, Frankliniella schultzei and Frankliniella occidentalis, in a persistant-propagative manner, and also by mechanical sap-inoculation. For more details of GRSV, refer to Arachis hypogaea.

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Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV occurs wherever Lactuca sativa crops are grown (Vicchi and Bellardi 1997; Vicchi et al. 1999; Koike et al. 2008; Kuo et al. 2014). The virus-infected lettuce plants develop brown to dark brown spots and dead (necrotic) areas. As necrosis spreads, much of the leaf browns dries out and dies. Leaf yellowing and brown spots are observed on both older and newer leaves. Margins of leaves may wilt and become yellow. Often only one side of the plant is affected. Plants infected early in development may become stunted and then die. Infected plants that survive to harvest are usually unmarketable. The virus is transmitted by the western flower thrips, Frankliniella occidentalis, in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of host plants. For more details of INSV, refer to Impatiens spp.

Lettuce big-vein associated varicosavirus Taxonomic position Genus: Varicosavirus

(LBVaV)

Family: Rhabdoviridae

Geographical distribution Lettuce big-vein disease was first described in plants of Lactuca sativa from California (Jagger and Chandler 1934). The virus spreads in Argentina, Brazil, Chile, Australia, the former Czechoslovakia, Slovenia, Italy, Iran, Saudi Arabia, Japan, the Netherlands, New Zealand, the UK, Mexico, and the USA (Colariccio et al. 2003; Roggero et al. 2003; Navarro et al. 2004; Rosales et al. 2004; Colariccio et al. 2005; Mavric Plesko et al. 2009; Tabarrozzi et al. 2010; Sorrentino et al. 2011; Alemzadeh and Izadpanah 2012; Verbeek et al. 2013; Ochoa-Martinez et al. 2014; Al-Saleh et al. 2015; Umar et al. 2017). Symptoms and host(s) Symptoms vary seasonally. Symptoms start as vein yellowing, and then develop to chlorotic or white vein banding. Often typical big-vein symptoms are produced (Colariccio, et al. 2003; Roggero et al., 2003). Infection frequently occurs in association with Mirafiori lettuce big-vein virus (MiLBVV), an Ophiovirus, which in single infection causes vein-band chlorosis; LBVaV in single infection induces necrotic spots and rings (Verbeek et al. 2013). The natural host range of this virus includes lettuce, endive (Cichorium endivia), spiny sow thistle (Sonchus asper), and common sow thistle (Sonchus oleraceus). Transmission The virus is transmitted by fungal vectors, Olpidium brassicae and O. virulentus (Lot et al. 2002; Maccarone 2009; Nomiyama et al. 2013). The virus is transmissible by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants and also not by seed or pollen (Huijberts et al. 1990). The resting spores of the fungus can persist for over 20 years in soil and can retain the ability to transmit the disease for over 15 years.

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Lactuca sativa (Lettuce)

Virion properties and genome The virions are rod-shaped and measures about 320–350 nm in length and 18 nm in width. The genome is bipartite, consisting of two negative-sense single-stranded RNAs: RNA 1 encodes the L protein, and is 6797 nt (AB075039 = NC_0011558) ; RNA2 encodes the coat protein and three other glycoproteins, and is 6081 nt (AB114138 = NC_011568) (Sasaya et al. 2002, 2004, 2008). This virus consists of a single CP with an Mr of 48 kDa and does not have an enveloped structure (Kuwata et al. 1983; Sasaya et al. 2011; Walker et al. 2018).

Lettuce chlorosis virus Taxonomic position Genus: Crinivirus

(LCV)

Family: Closteroviridae

Geographical distribution LCV infection in plants of Lactuca sativa occurs in deserts of California and Southwestern USA (Duffus et al. 1996; McLain et al. 1998; Salem et al. 2009). Symptoms and host(s) The virus-infected lettuce plants show severe yellowing and/or reddening and stunting, rolling, vein-clearing, and brittleness of leaves. The virus has a wide host range including lettuce, sugar beet, and several other crops and weeds (but not among the Cucurbitaceae). Transmission The virus is transmitted by B. tabaci biotypes A and B with similar efficiencies in a semi-persistent manner. Whiteflies can acquire and transmit the virus with AAP/IAP of 1 h each (Duffus et al. 1996). Transmission was more efficient after 24 h of feeding, whereas retention did not exceed 4 days. The virus is not mechanically sap-transmissible and also not through seed. Virion properties and genome The virions are non-enveloped, bipartite filaments particles about 650–850 nm and 700–900 nm in length, and 10–13 nm in diameter. The genome consists of two molecules of positive-sense ssRNA. RNA1 consists of 8591 nt (FJ380118 = NC_012909) and RNA2 consists of 8556 nt (FJ380119 = NC_012910) (Salem et al. 2009; Liu et al. 2000; Kreuze 2011; Tzanetakis et al. 2013; Zhao et al. 2017).

Lettuce infectious yellows virus Taxonomic position Genus: Crinivirus

(LIYV)

Family: Closteroviridae

Geographical distribution LIYV infection in plants of Lactuca sativa occurs in North America and Mexico (Duffus et al. 1982; Brown and Stanghellini 1988; Brown and Poulos 1989; Brown et al. 1990; Rubio et al. 1999; Tzanetakis et al. 2013; Abrahamian and Abou-Jawdah 2014).

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Symptoms and host(s) The virus-infected lettuce plants cause interveinal chlorosis which leads to severe leaf yellowing, rolling, and brittleness of leaves. Infected plants are usually stunted and, in head lettuce cultivars, compact heads fail to develop. Necrotic lesions appear at or near margins of older leaves as the disease progresses (Brown and Nelson 1986). LIYV has a wide host range (45 species in 15 families) and infects economically important crops like beetroot (Beta vulgaris), lettuce (Lactuca sativa), marrows (Cucurbita pepo), melons (Cucumis melo), carrots (Daucus carota), cucurbits (Cucurbita foetidissima, C. maxima, C. moschata), and watermelons (Citrullus lanatus) (Halliwell and Johnson 1992). Transmission The virus is transmitted by the whitefly vector B. tabaci biotype B in a semi-persistent manner, (Duffus et al. 1986; Cohen et al. 1992; Jones 2003; Gilbertson et al. 2015). It is retained by viruliferous whiteflies for several days in serial transfers on susceptible hosts (Duffus et al. 1986). The closely related whitefly B. argentifolii is a very inefficient vector. The virus is not mechanically saptransmissible and no seed-transmission is reported. Virion properties and genome The virions are non-enveloped, bipartite filaments particles about 650–850 nm and 700–900 nm in length, and 10–13 nm in diameter. The genome consists of two molecules of positive-sense ssRNA. RNA1 consists of 8118 nt (U15440 = NC_003617) and RNA2 of 7193 nt (U15441 = NC_003618) (Larsen et al. 1988; Klaassen et al. 1995; Tian et al. 1999; Kreuze 2011).

Lettuce Italian necrotic virus (LINV) Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution LINV infection in plants of Lactuca sativa was reported from Italy and southeastern France (Ciuffo et al. 2016; Desbiez et al. 2017). Symptoms and host(s) The virus-infected lettuce plants exhibit necrosis and leaf distortion symptoms. Transmission The virus is transmitted by an aphid vector Myzus persicae in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, and 680–900 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9829 nt (KP769852 = NC_027706) (Revers and Garcia 2015; Wylie et al. 2017).

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Lettuce mosaic virus Taxonomic position Genus: Potyvirus

Lactuca sativa (Lettuce)

(LMV)

Family: Potyviridae

Geographical distribution LMV was first reported in plants of Lactuca sativa from Florida in 1921, and since then this virus has been reported from almost all lettuce-growing areas in the world (Al-Musa and Mansour 1985; Lot and Maury-Chovelon 1985; Alhubaishi et al. 1987; Lot 1988; Chen et al. 1995; Twardowicz-Jakusz and Zielinska 1997; Stangarlin et al. 2000; Moreno et al. 2004, 2007; Firmino et al. 2008; Pavan et al. 2008; Al-Saleh et al. 2009; Soleimani et al. 2011; Sharma et al. 2013). Symptoms and host(s) The virus-infected lettuce plants generally exhibit mosaic and crinkling of the lamina and blistering of leaves. At the very early stage of virus infection young plants are stunted, deformed, and rarely grow to full size; head lettuce varieties fail to form heads. Plants that are infected later in the growth cycle will show a different set of symptoms. These plants may reach full size, but the older outer leaves will be yellow, twisted, and otherwise deformed. On head lettuce the wrapper leaves often will curve back away from the head. Developing heads may be deformed. In some cases brown, necrotic flecks occur on the wrapper leaves (Krause-Sakate et al. 2004). This virus has a wide natural host range; susceptible species have been identified in 20 genera belonging to 10 different families including Compositae, Chenopodiaceae, and Cruciferae (Dinant and Lot 1992; Abdel Wahab 2012). Transmission The virus is transmitted by aphid vectors including Aphis gossypii, Macrosiphum euphorbiae, and Myzus persicae, in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and has a very wide host range. Seed-transmission up to 15% in lettuce was recorded (Grogan et al. 1952), and even very low rates of seed-transmission may result in significant crop losses. The virus is also transmitted by pollen to the seed. Virion properties and genome The virions are non-enveloped, flexuous filaments, with a clear modal length of 750 nm long and 15 nm wide. The genome consists of a single molecule of linear, positive-sense single-stranded RNA of 10,080 nt (X97705 = NC_003605) (Tomlinson 1964; Revers et al. 1997, 1999; German-Retana et al. 2008; Sharma et al. 2014; Revers and Garcia 2015; Wylie et al. 2017).

Lettuce necrotic leaf curl virus Taxonomic position Genus: Torradovirus

(LNLCV)

Family: Secoviridae

Geographical distribution The virus infection in plants of Lactuca sativa was reported from the Netherlands (Verbeek et al. 2014; van der Vlugt et al. 2015).

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Symptoms and host(s) The virus-infected lettuce plants exhibit leaf necrosis and curling symptoms. Transmission The virus is transmissible by mechanical sap-inoculation to different lettuce cultivars and Nicotiana species. Virion properties and genome The virions are isometric, non-enveloped of two types, but similar in size 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is a bipartite, linear positive-sense, single-stranded RNA. RNA1 (7586 nt; KC855266 = NC_035214) consists of a single open reading frame. The RNA1-ORF1 of 6669 nt encodes a predicted polyprotein of 2223 amino acids (aa) with a molecular mass of 246 kDa. RNA2 (5300 nt; KC855267 = NC_035219) contains two open reading frames; RNA2-ORF1 encodes a predicted protein of 211 aa (23 kDa), and RNA2-ORF2 encodes for a predicted polyprotein of 1222 aa (134 kDa) (Verbeek et al. 2014; Sanfacon 2015; van der Vlugt et al. 2015; Thompson et al. 2017).

Lettuce necrotic yellows cytorhabdovirus Taxonomic position Genus: Cytorhabdovirus

(LNYV)

Family: Rhabdoviridae

Geographical distribution LNYV is often severe in Lactuca sativa crops in Australia and also occurs in New Zealand but is of little or no importance in other countries (Stubbs and Grogan 1963; Francki and Randles 1970; Fry et al. 1973; Randles 1993; Dietzgen et al. 2007; Higgins et al. 2016). Symptoms and host(s) The initial symptoms of virus-infected lettuce plants are the browning of leaf veins, followed by partial death of the inner leaves. Affected plants are yellow and stunted, often with twisted and lopsided leaves. In advanced stages, the outer leaves wilt severely, giving the plant a flattened, stunted appearance. Transmission The virus is transmitted mainly by the sow thistle aphid, Hyperomyzus lactucae, in a persistentpropagative manner. H. carduellinus is a less important vector. The virus is also sap-transmissible and has a narrow host range in the Chenopodiaceae, Compositae, Leguminosae, Liliaceae, and Solanaceae. The virus is not transmitted by seed. Virion properties and genome The virions are enveloped, bacilliforms and measure 52 nm in diameter and 360 nm long. There are distinct surface projections (spikes) dispersed evenly over all the surface. The genome is a monopartite, negative-sense, single-stranded RNA of 12,807 nt (AJ867584 = NC_007642) is unsegmented (Wetzel et al. 1994; Callaghan and Dietzgen 2005; Dietzgen et al. 2006, 2007; Dietzgen 2011; Walker et al. 2018).

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Lactuca sativa (Lettuce)

Lettuce ring necrosis ophiovirus Taxonomic position Genus: Ophiovirus

(LRNV)

Family: Aspiviridae

Geographic distribution LRNV infection in plants of Lactuca sativa is known to occur in Belgium, the Netherlands, France, and the USA (Bos and Huijberts 1996; Campbell and Lot 1996). Symptoms and host(s) The virus induces severe disease symptoms on butterhead and crisphead lettuce whereas mild symptoms occur on iceberg and cos lettuce. Butterhead lettuce exhibits yellow spots giving an irregular blotchy appearance on the basal and intermediate leaves, and necrotic rings develop on the midribs. Necrotic rings and yellow mottling develop on the basal leaves of crisphead lettuce. Transmission The virus is transmitted through a fungal vector, Olpidium brassicae (Campbell and Lot, 1996). The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, naked filamentous nucleocapsids about 3 nm in diameter, forming kinked circles of at least two different contour lengths (300–500 and 1500–2500 nm). The circles (open form) can collapse to form pseudo-linear duplex structures of 9–10 nm in diameter (collapsed form). The genome is a linear, negative-sense, single-stranded RNA and consists of four segments. RNA1 consists of 7651 nt (AY535016 = NC_006051), RNA2 of 1830 nt (AY535017 = NC_006052), RNA3 of 1527 nt (AY535018 = NC_006053), and RNA4 of 1417 nt (AY535019 = NC_006054) (Milne et al. 2011; Garcia 2012).

Lettuce speckles mottle virus Taxonomic position Genus: Umbravirus

(LSMV)

Family: Tombusviridae

Geographical distribution LSMV infection was first reported in plants of Lactuca sativa from the USA (Falk et al. 1978, 1979a, b). The virus spreads in the USA (California). Symptoms and host(s) LSMV in association with Beet western yellows virus causes speckles disease of lettuce, which is characterized by small angular chlorotic spots on the outer leaves of the infected plants. The disease symptom is severe during early spring, and it disappears with the onset of summer (Falk et al. 1979b). Transmission The virus is transmitted by aphid vectors, Acyrthosiphon solani, Brevicoryne brassicae, and Myzus persicae, in a circulative, non-propagative manner. LSMV requires a helper virus (Beet western yellows luteovirus) for vector transmission. The virus is transmissible by mechanical sap-inoculation to several three to nine families which are susceptible.

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Virion properties and genome The virus does not form typical virus particles as it lacks a functional coat protein; however, 50–70 nm spherical membranous structures are associated with the tonoplast in vacuoles of infected cells (Falk et al. 1979a). The genome is a positive-sense ssRNA. Genome sequence information is not available. The infected lettuce plant tissues contain abundant double-stranded RNA (ds-RNA) and ss-RNA, and only the ssRNA was infectious (Falk et al. 1979a).

Lettuce virus X

(LeVX)

Taxonomic position Genus: Potexvirus

Family: Alphaflexiviridae

Geographical distribution LeVX infection in plants of Lactuca sativa was reported from Iran (Dizadji et al. 2008). Symptoms and host(s) The virus-infected lettuce plants are generally symptomless even though they contain high concentration of the virus. Transmission The virus is mechanically sap-transmissible. The virus causes systemic vein yellowing in the assay host Chenopodium quinoa and can also infect Nicotiana benthamiana. Virion properties and genome The virions are flexuous filaments 470–580 nm in length and 13 nm in diameter, with a helical symmetry and a pitch of 3.3–3.7 nm. The genome consists of a single linear molecule of positivesense ssRNA of 7212 nt (AM745758 = NC_010832) and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type and 18–27 kDa in size (Adams et al. 2004; Dizadji et al. 2008).

Lettuce yellow mottle cytorhabdovirus Taxonomic position Genus: Cytorhabdovirus

(LYMoV)

Family: Rhabdoviridae

Geographical distribution LYMoV infection in plants of Lactuca sativa was reported from France and South America (Jadao et al. 2007; Heim et al. 2008). Symptoms and host(s) The virus-infected lettuce plants exhibit typical chlorotic spots and yellow leaf mottling symptoms on leaves. Virion properties and genome The virions are enveloped, bacilliforms and measure 60–75 nm in diameter and 200–350 nm long. There are distinct surface projections (spikes) dispersed evenly over all the surface. The genome is a

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Lactuca sativa (Lettuce)

monopartite, negative-sense, single-stranded RNA of 12,926 nt (EF687738 = NC_011532) and contains six open reading frames (Heim et al. 2008; Dietzgen 2011; Walker et al. 2018).

Lettuce yellows virus

(LYV)

Taxonomic position LYV is a tentative member of the genus Polerovirus and family Luteoviridae. Geographical distribution LYV infection in plants of Lactuca sativa was reported from Greece (Lotos et al. 2016). Symptoms and host(s) The virus-infected lettuce plants exhibit mild to severe interveinal yellowing symptoms. Transmission The virus is transmitted by aphid vectors in a persistent manner. Virion properties and genome The virions are 25–30 nm in diameter. The genome is a positive-sense single-stranded RNA; only partial sequences are currently available (LN865080, LT595023).

Mirafiori lettuce big-vein ophiovirus Taxonomic position Genus: Ophiovirus

(MiLBVV)

Family: Aspiviridae

Geographical distribution MiLBVV infection in plants of Lactuca sativa occurs in the USA, Brazil, Japan, Chile, Argentina, Iran, Europe, and Mediterranean Region (Rosales et al. 2004; Navarro et al. 2004; Colariccio et al. 2005; Sasaya et al. 2008; Mavric Plesko et al. 2009; Maccarone et al. 2010; Tabarrozzi et al. 2010; Sorrentino et al. 2011; Alemzadeh and Izadpanah 2012; Al-Saleh et al. 2015). Symptoms and host(s) The virus-infected lettuce plants cause the veins in leaves to become very enlarged and clear. Such deformities are easily seen if the leaf is examined with the light source behind it. The enlarged veins cause the rest of the leaf to be ruffled, twisted, and malformed. Severely affected plants can be so deformed as to be unmarketable, and head lettuce varieties may fail to form a head. However, infected plants showing less severe symptoms can still be harvested (Natsuaki et al. 2002; Roggero et al. 2003). Transmission This virus disease is soil-borne, and fungal vectors, Olpidium brassicae and O. virulentus, transmit this virus (Lot et al. 2002; Maccarone 2009; Nomiyama et al. 2013). Primary infections occur via the zoospores which are released by the resting spots. The mobile spores infect the epidermal cells of the young roots of lettuce. The fungus remains on the roots and forms numerous zoosporangia that produce a large number of zoospores, some of which become viruliferous. The virus is mechanically

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sap-transmissible, and species belonging to the families Chenopodiaceae, Compositae, Solanaceae, and Tetragoniaceae are susceptible. Virion properties and genome The virions are non-enveloped, naked filamentous nucleocapsids about 3 nm in diameter, forming kinked circles of at least two different contour lengths (300–500 and 1500–2500 nm). The circles (open form) can collapse to form pseudo-linear duplex structures of 9–10 nm in diameter (collapsed form). The genome is a linear, negative-sense, single-stranded RNA and consists of four segments. RNA1 consists of 7794 nt (AF525933), RNA2 of 1788 nt (AF525934), RNA3 of 1515 nt (AF525935), and RNA4 of 1402 nt (AF525936) (Lot et al. 2002; van der Wilk et al. 2002; Kawazu et al. 2003; Milne et al. 2011; Garcia 2012).

Moroccan pepper virus (MPV) Synonyms Lettuce necrotic stunt virus (LNSV) Taxonomic position Genus: Tombusvirus

Family: Tombusviridae

MPV (initially reported as LNSV) infection in plants of Lactuca sativa was reported from the southwestern USA (Obermeier et al. 2001; Wintermantel and Anchieta 2012; Wintermantel and Hladky 2013). The virus-infected lettuce plants are severely stunted and mature, diseased plants may only reach 6–8 in. in height. The outermost leaves are extensively yellowed. The younger, inner leaves often remain dark green in color, but rough and leathery in texture. In some cases, the older leaves develop necrotic spotting that can turn into extensive areas of brown, dead tissue. The virus is mechanically saptransmissible. The virus is soil-borne and no biological vector is known. The virus is spread through contaminated soil and water. For more details of MPV, refer to Capsicum anuum.

Sonchus yellow net nucleorhabdovirus Taxonomic position Genus: Nucleorhabdovirus

(SYNV)

Family: Rhabdoviridae

SYNV was reported to infect plants of Lactuca sativa in Florida, USA (Falk et al. 1986). The virusinfected lettuce plants exhibit bright yellow interveinal spotting on older leaves. The virus is transmitted by aphid vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of SYNV, refer to Lactuca sativa.

Sowthistle yellow vein nucleorhabdovirus Taxonomic position Genus: Nucleorhabdovirus

Family: Rhabdoviridae

(SYVV)

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Lactuca sativa (Lettuce)

SYVV was identified as infecting plants of Lactuca sativa in the USA (Duffus et al. 1970). The virusinfected lettuce plants exhibit symptoms of vein-clearing, vein yellowing, and stunting. The virus is transmitted by an aphid vector, Hyperomyzus lactucae, in a persistent, propagative manner. The virus is not transmissible by mechanical sap-inoculation, and also not by contact between plants. For more details of SYVV, refer to Sonchus spp.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV was reported to infect plants of romaine lettuce in the USA (Mayhew and Matsumoto 1978). The virus-infected lettuce plants exhibit stunted and flattened growth, a brilliant, chrome-yellow ring-spot, and line patterns on the leaves. A nematode (Paratrichodorus christei) is the vector of the virus. The virus is transmissible by mechanical sap-inoculation to a large number of host plants. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Lactuca sativa was reported from Iran (Abtahi and Koohi Habibi 2008; Abtahi and Khodai Motlagh 2009). The virus-infected lettuce plants show symptoms such as mosaic, vein-clearing, vein necrosis, yellowing, and leaf distortion; necrosis of petioles and stems and death of the plants may follow. This virus is transmitted by thrips vectors, Frankliniella schultzei and Thrips tabaci; the virus present in/on pollen entering the host through injuries caused by thrips while feeding. The virus is mechanically sap-transmissible to a number of hosts. The virus is pollen transmitted. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato bushy stunt virus Taxonomic position Genus: Tombusvirus

(TBSV)

Family: Tombusviridae

TBSV infection in plants of Lactuca sativa was reported from California, USA (Novak et al. 1981; Liu et al. 1999). The virus-infected lettuce plants exhibit dieback and necrotic symptoms. The virus is transmissible by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants. There is no known vector for this virus. For more details of TBSV, refer to Solanum lycopersicum.

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Tomato chlorotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(TCSV)

Family: Tospoviridae

TCSV infection in plants of Lactuca sativa occurs in Argentina, Brazil, and Puerto Rico (Gracia et al. 1999; Dal Bo et al. 2001; Colariccio et al. 2001, 2004; Pavan et al. 2008; de Jensen and Adkins 2014). The affected lettuce plants exhibit symptoms including dwarfing, mosaic, chlorotic and necrotic ringspots, and necrosis. The virus is transmitted by thrips vectors, Frankliniella schultzei and Frankliniella occidentalis, in a persistent propagative manner. The virus is also mechanically saptransmissible. For more details of TCSV, refer to Solanum lycopersicum.

Tomato infectious chlorosis virus Taxonomic position Genus: Crinivirus

(TICV)

Family: Closteroviridae

TICV infection in plants of Lactuca sativa was reported from Italy (Parrella and Filella 2007; Parrella 2008). The virus-infected lettuce plants exhibit symptoms of interveinal yellowing on the leaves. The virus is transmitted by the whitefly vector, Trialeurodes vaporariorum, in a semi-persistent manner. The virus is not transmissible by mechanical inoculation. For more details of TICV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Lactuca sativa was reported from Mediterranean/Middle Eastern countries (Israel, Jordan, Turkey,Iran, Saudi Arabia, Spain), Brazil, and Hawaii (Cho et al. 1987; Wilson 1998; Gera et al. 2000; Moreno et al. 2004; Pavan et al. 2008; Kamberoglu and Alan 2011; Soleimani et al. 2011; Salem et al. 2012; Al-Saleh et al. 2014). The virus-infected lettuce plants show necrotic lesions, varying from light brown to black on young leaves and petioles. Old leaves display chlorotic spots which become necrotic at a later stage. The discoloration extends to the heart leaves, and cessation of growth on one side of the plant produces characteristic symptoms. In some cases plants grow slowly, are stunted, and finally die. If they survive, their heads fail to form. The virus is transmitted by a thrips vector, F. occidentalis, in a persistent propagative manner, and also by mechanical sap-inoculation (Cho et al. 1989). The virus has a very wide host range. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Turnip curly top virus Taxonomic position Genus: Turncurtovirus

(TCTV)

Family: Geminiviridae

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Lactuca sativa (Lettuce)

TCTV infection in plants of Lactuca sativa was reported from Iran (Kamali et al. 2016). The virus is transmitted by the leafhopper vector Circulifer haematoceps in a circulative and non-propagative manner. For more details of TCTV refer to Brassica rapa.

Turnip leaf roll virus Taxonomic position Genus: Turncurtovirus

(TuLRV)

Family: Geminiviridae

TuLRV infection in plants of Lactuca sativa was reported from Iran (Kamali et al. 2016). The virusinfected lettuce plants exhibit dark green vein banding symptoms. The virus is transmitted by the leafhopper vector Circulifer haematoceps in a circulative, non-propagative manner. For more details of TuLRV refer to Brassica rapa.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Lactuca sativa was reported from Europe and the USA (Moreno et al. 2004; Blancard et al. 2006; Sanchez et al. 2007). The virus-infected lettuce plants exhibit early symptoms which include numerous, small, light green, circular, and irregular lesions on leaves. Later symptoms include curvature of the midrib and asymmetrical distortion of the leaf blade. The lamina often becomes necrotic; unlike lettuce infected with beet western yellows virus, veins do not become green, and leaves do not become brittle in TUMV-infected plants. The oldest leaves often become bright yellow all over. Lettuce infected early may be stunted. The disease is occasionally lethal. The virus is transmissible by many species of aphids in a non-persistent manner. The virus is transmitted by mechanical sap-inoculation, and has a wide host range in 20 plant families and is especially common in crucifers. The virus is not seed-transmitted in lettuce. For more details of TuMV, refer to Brassica rapa.

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Lactuca sativa (Lettuce)

Natsuaki KT, Morikawa T, Natsuaki T, Okuda S (2002) Mirafiori lettuce virus detected from lettuce with big vein symptoms in Japan. Jpn J Phytopathol 68:309–312 Navarro JA, Botella F, Maruhenda A, Sastre P, Sanchez-Pina MA, Pallas V (2004) Comparative infection progress analysis of Lettuce big-vein virus and Mirafiori lettuce virus in lettuce crops by developed molecular diagnosis techniques. Phytopathology 94:470–477 Nomiyama K; Inoue H; Sekiguchi H; Tomioka K; Osaki H; Takehara T; Sasaya T (2013) Transmission of lettuce big-vein associated virus and Mirafiori lettuce big-vein virus by Olpidium brassicae. In: Proceedings of the ninth symposium of the international working group on plant viruses with fungal vectors, Obihiro, Hokkaido, 19–22 Aug 2013, pp 25–27 Novak JB, Novakova J, Lanzova J (1981) Demonstration of tomato bushy stunt virus in lettuce. Ochr Rostl 17:241 Obermeier C, Sears JL, Liu HY, Schluenter KO, Ryder EJ, Duffus JE, Koike ST, Wisler GC (2001) Characterization of distinct tombusviruses that cause diseases of lettuce and tomato in the western United States. Phytopathology 91:797–806 Ochoa-Martinez DL, Alfonsina-Hernandez J, Sanchez-Escudero J, Rodriguez-Martinez D, Vera-Graziano J (2014) First report of Lettuce big-vein associated virus (Varicosavirus) infecting lettuce in Mexico. Plant Dis 98:573 Parrella G (2008) Interveinal yellowing caused by Tomato infectious chlorosis virus in lettuce and escarole in southern Italy. J Phytopathol 156:190–192 Parrella G, Filella F (2007) A severe outbreak of Tomato infectious chlorosis virus in lettuce and escarole in Southern Italy. J Plant Pathol 89(3):S52 Pavan MA, Krause-Sakate R, da Silva N, Zerbini FM, Le Gall O (2008) Virus diseases of lettuce in Brazil. Plant Viruses 2:35–41 Purcifull DE, Christie SR, Zitter TA, Bassett MJ (1971) Natural infection of lettuce and endive by Bidens mottle virus. Plant Dis Reptr 55:1061–1063 Randles JW (1993) Plant viruses, viroids and virologists of Australasia. Aust Plant Pathol 22(4):122–130 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Revers F, Yang SJ, Walter J, Souche S, Lot H, Le Gall O, Candresse T, Dunez J (1997) Comparison of the complete nucleotide sequences of two isolates of lettuce mosaic virus differing in their biological properties. Virus Res 47(2):167–177 Revers F, van der Vlugt RAA, Souche S, Lanneau M, Lot H, Candresse T, Le Gall O (1999) Nucleotide sequence of the 30 -terminal region of the genome of four lettuce mosaic virus isolates from Greece and Yemen. Arch Virol 144:1619–1626 Roggero P, Lot H, Souche S, Lenzi R, Milne RG (2003) Occurrence of Mirafiori lettuce virus and Lettuce big-vein virus in relation to development of big-vein symptoms in lettuce crops. Eur J Plant Pathol 109:261–267 Rosales M, Sepulveda P, Bruna A (2004) First report of Lettuce big-vein virus and Mirafiori lettuce virus in Chile. Plant Dis 88:1286 Rubio L, Soong J, Kao J, Falk BW (1999) Geographic distribution and molecular variation of isolates of 3 whitefly-borne closteroviruses: of cucurbits, Lettuce infectious yellows virus, Cucurbit yellow stunting disorder virus and Beet pseudo yellows virus. Phytopathology 89:707–711 Salem NM, Chen AY, Tzanetakis IE, Mongkolsiriwattana C, Ng JC (2009) Further complexity of the genus Crinivirus revealed by the complete genome sequence of Lettuce chlorosis virus (LCV) and the similar temporal accumulation of LCV genomic RNAs 1 and 2. Virology 390:45–55 Salem NM, Mansour A, Badwan H (2012) Identification and partial characterization of Tomato spotted wilt virus on lettuce in Jordan. J Plant Pathol 94:431–435 Sanchez F, Rodriguez-Mateos M, Tourino A, Fresno J, Gomez-Campo C, Jenner CE, Walsh JA, Ponz F (2007) Identification of new isolates of Turnip mosaic virus that cluster with less common viral strains. Arch Virol 152:1061–1068 Sanfacon H (2015) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. In: Encyclopedia of Life Sciences. Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000764.pub3. http:// www.els.net Sasaya T, Ishikawa K, Kogenezawa H (2002) The nucleotide sequence of RNA1 of Lettuce big vein virus, genus Varicosavirus reveals it relation to non-segmented negative stranded RNA viruses. Virology 297:289–297 Sasaya T, Kusaba S, Ishikawa K, Koganezawa H (2004) Nucleotide sequence of RNA2 of Lettuce big-vein virus and evidence for a possible transcription termination/initiation strategy similar to that of rhabdoviruses. J Gen Virol 85(PT 9):2709–2717 Sasaya T, Fujii H, Ishikawa K, Koganezawa H (2008) Further evidence of Mirafiori lettuce big-vein virus but not of Lettuce big-vein associated virus with big-vein disease in lettuce. Phytopathology 98:464–468 Sasaya T, Milne RG, Campbell RN, Josef Vetten H (2011) Varicosavirus. Unassigned genus. In: The Springer Index of Viruses. Springer, New York, pp 2081–2085. https://doi.org/10.1007/978-0-387-95919-1_340 Sharma P, Jain RK, Saha S, Kalia P (2013) First report of Lettuce mosaic virus infecting Lactuca sativa in India. Plant Dis 97:849

Lactuca sativa (Lettuce)

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Sharma P, Sahu AK, Verma RK, Mishra R, Choudhary DK, Gaur RK (2014) Current status of Potyvirus in India. Arch Phytopathol Plant Prot 47:906–918 Soleimani P, Mosahebi G, Habibi MK (2011) Identification of some viruses causing mosaic on lettuce and characterization of Lettuce mosaic virus from Tehran Province in Iran. Afr J Agric Res 6:3029–3035 Sorrentino R, Carrieri R, Cirillo G, Parrella G, Alioto D (2011) Severe outbreaks of lettuce big vein in Latium and Campania. J Plant Pathol 93:S4.57 Stangarlin O, Pavan MA, da Silva N (2000) Occurrence of a new pathotype of Lettuce mosaic virus on lettuce in Brazil. Plant Dis 84(4):490 Stone WJH, Nelson MR (1966) Alfalfa mosaic virus (Calico) of lettuce. Plant Dis Reptr 50:629–631 Stubbs LL, Grogan RG (1963) Necrotic yellows: a newly recognized virus disease of lettuce. Aust J Agric Res 14:439–459 Suzuki GS, Rosa RAC, Sanches MM, Nozaki DN, Pavan MA, Krause-Sakate R (2009) Characterization of an isolate of Bidens mosaic virus (BiMV) from lettuce. Summa Phytopathol 35:231–233 Tabarrozzi AEB, Pena EJ, Dal Bo E, Robles Luna G, Reyes CA, Garcia ML (2010) Identification of Mirafiori lettuce bigvein virus and Lettuce big-vein associated virus infecting Lactuca sativa with symptoms of lettuce big-vein disease in Argentina. Plant Pathol 59:1160–1161 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J Gen Virol 98:529–531 Tian T, Rubio L, Yeh HH, Crawford B, Falk BW (1999) Lettuce infectious yellows virus: in vitro acquisition analysis using partially purified virions and the whitefly, Bemisia tabaci. J Gen Virol 80:1111–1117 Timmerman EL, D’Arcy CJ, Splittstoesser WE (1985) Beet western yellows virus in Illinois vegetable crops and weeds. Plant Dis 69:933–936 Tomlinson JA (1964) Purification and properties of lettuce mosaic virus. Ann Appl Biol 53:95–102 Tomlinson JA, Walker VM (1973) Beet western yellows virus in lettuce. Report of the National Vegetable Research Station for 1972, pp 97–98 Twardowicz-Jakusz A, Zielinska L (1997) Occurrence and identification of viruses affecting lettuce in Poland. I. Survey of viruses affecting lettuce field plantations. J Plant Prot Res 37:11–17 Tzanetakis IE, Martin RR, Wintermantel WM (2013) Epidemiology of Criniviruses: an emerging problem in world agriculture. Front Microbiol 4:119. https://doi.org/10.3389/fmicb.2013.00119 Umar M, Amer MA, Al-Saleh MA, Al-Shahwan IM, Shakeel MT, Zakri AM, Katis NI (2017) Characterization of lettuce bigvein associated virus and Mirafiori lettuce big-vein virus infecting lettuce in Saudi Arabia. Arch Virol 162:2067–2072 van der Vlugt RAA, Verbeek M, Dullemans AM, Wintermantel WM, Cuellar WJ, Fox A, Thompson JR (2015) Torradoviruses. Annu Rev Phytopathol 53:485–512 van der Wilk F, Dullemans AM, Verbeek M, van den Heuvel JFJM (2002) Nucleotide sequence and genomic organization of an Ophiovirus associated with lettuce big vein disease. J Gen Virol 83:2869–2877 Verbeek M, Dullemans AM, van Bekkum PJ, van der Vlugt RAA (2013) Evidence for Lettuce big-vein associated virus as the causal agent of a syndrome of necrotic rings and spots in lettuce. Plant Pathol 62:444–451 Verbeek M, Dullemans AM, van Raaij HMG, Verhoeven JTJ, van der Vlugt RAA (2014) Lettuce necrotic leaf curl virus, a new plant virus infecting lettuce and a proposed member of the genus Torradovirus. Arch Virol 159:801–805 Vetten HJ, Breyel E, Lesemann DE, Maiss E, Weidemann HL (1985) Properties of an isometric lettuce virus resembling Dandelion yellow mosaic virus. Phytoparasitica 13:3–4 Vicchi V, Bellardi MG (1997) Impatiens necrotic spot tospovirus infection on lettuce in Italy. Informatore Fitopatologico 47:55–57 Vicchi V, Fini P, Cardoni M (1999) Presence of Impatiens necrotic spot tospovirus (INSV) on vegetable crops in EmiliaRomagna region. Informatore Fitopatologico 49(4):53–55 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Rhabdoviridae. J Gen Virol 99:447–448 Wang HY, Li XD, Liu YY, Wang B, Zhu XP (2008) First report of Beet mosaic virus infecting lettuce, in China. Plant Pathol 57:764 Wetzel T, Dietzgen RG, Dale JL (1994) Genomic organization of lettuce necrotic yellows rhabdovirus. Virology 200:401–412 Wilson CR (1998) Incidence of weed reservoirs and vectors of tomato spotted wilt tospovirus on southern Tasmanian lettuce farms. Plant Pathol 47:171–176 Wintermantel WM, Anchieta AG (2012) The genome sequence of lettuce necrotic stunt virus indicates a close relationship to Moroccan pepper virus. Arch Virol 157:1407–1409. https://doi.org/10.1007/s00705-012-1307-x Wintermantel WM, Hladky LL (2013) Complete genome sequence and biological characterization of Moroccan pepper virus (MPV) and reclassification of Lettuce necrotic stunt virus as MPV. Phytopathology 103:501–508 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

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Lactuca sativa var. augustana (Celtuce plant)

Xiang HY, Shang QX, Han CG, Li DW, Yu JL (2008) First identification of Beet western yellows virus on sugarbeet and lettuce in China. Plant Pathol 57:390 Zhao X, Zhu M, Wu Q, Zhang J, Xu Y, Tao X (2017) Complete genome sequence of a lettuce chlorosis virus isolate from China and genome recombination/rearrangement analysis. Arch Virol 163:751–754 Zink FW, Duffus JE (1972) Association of Beet western yellows and Lettuce mosaic viruses with internal rib necrosis of lettuce. Phytopathology 62:1141–1144

Lactuca sativa var. augustana (Celtuce plant) Family: Asteraceae

Leafy vegetable

Calla lily chlorotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(CCSV)

Family: Tospoviridae

CCSV infection in plants of Lactuca sativa var. augustana was reported from China (Wu et al. 2018). The virus-infected celtuce plants exhibit vein-clearing and chlorotic spot symptoms. The virus is transmitted by thrips vectors in a persistent propagative manner, and also by mechanical sap-inoculation. For more details of CCSV, refer to Zantedeschia spp.

References Wu X, Wu X, Li W, Cheng X (2018) Molecular characterization of a divergent strain of calla lily chlorotic spot virus infecting celtuce (Lactuca sativa var. augustana) in China. Arch Virol 163:1375–1378

Lactuca serriola (Prickly lettuce) Family: Asteraceae

Beet curly top virus Taxonomic position Genus: Curtovirus

Weed host

(BCTV)

Family: Geminiviridae

BCTV infection in plants of Lactuca serriola was reported from California, USA (Creamer et al. 1996). The virus is transmitted by a leafhopper vector, Circulifer tenellus, in a persistent (circulative, non-propagative) manner. The virus is not transmissible by mechanical inoculation. For more details of BCTV, refer to Beta vulgaris.

Lactuca serriola (Prickly lettuce)

Lettuce mosaic virus Taxonomic position Genus: Potyvirus

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(LMV)

Family: Potyviridae

LMV infection in plants of Lactuca serriola was reported from France (German-Retana et al. 2008). The virus-infected prickly lettuce plants exhibit mosaic mottling symptoms on the leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is transmitted through seeds of L. serriola. For more details of LMV, refer to Lactuca sativa.

Plum pox virus Taxonomic position Genus: Potyvirus

(PPV)

Family: Potyviridae

PPV infection in plants of Lactuca serriola was reported from Bulgaria (Milusheva and Rankova 2002). The virus-infected prickly lettuce plants exhibit pale or yellow green ring-spots or mottling on leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PPV, refer to Prunus domestica.

L Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Lactuca serriola was reported from Australia (Sharman et al. 2015). The virus-infected prickly lettuce plants exhibit chlorosis symptoms. The virus is transmitted by thrips vectors; the virus present in/on pollen entering the host through injuries caused by thrips while feeding. The virus is mechanically sap-transmissible and not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Lactuca serriola was reported from the USA (Groves et al. 2002). The virus-infected prickly lettuce plants exhibit yellow spots or ring-spots on the leaves. The virus is transmitted by the thrips vector, Frankliniella fusca, in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

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Lagenaria breviflora

References Creamer R, Lugue-Williams M, Howo M (1996) Epidemiology and incidence of Beet curly top geminivirus in naturally infected weed hosts. Plant Dis 80:533–535 German-Retana S, Walter J, Le Gall O (2008) Lettuce mosaic virus: from pathogen diversity to host interactors. Mol Plant Pathol 9:127–136 Groves RL, Walgenbach JF, Mayor JW, Kennedy GG (2002) The role of weed hosts and tobacco thrips, Frankliniella fusca, in the epidemiology of Tomato spotted wilt virus. Plant Dis 86:573–582 Milusheva S, Rankova Z (2002) Plum pox potyvirus detection in weed species under field conditions. Acta Hortic 577:283–287 Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct Tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207

Lagenaria breviflora Family: Cucurbitaceae

Medicinal

Moroccan watermelon mosaic virus Taxonomic position Genus: Potyvirus

(MWMV)

Family: Potyviridae

MWMV infection in plants of Lagenaria breviflora was reported from Southeastern Nigeria (Owolabi et al. 2012). The virus-infected Lagenaria breviflora plants exhibit mosaic, leaf malformation and conspicuous green vein-banding symptoms. The virus is transmitted by aphid vectors in a nonpersistent manner, and also by mechanical sap-inoculation. For more details of MWMV, refer to Citrullus lanatus.

References Owolabi AT, Rabenstein F, Ehrig F, Maiss Edgar M, Vetten HJ (2012) Strains of Moroccan watermelon mosaic virus isolated from Lagenaria breviflorus and Coccinia barteri in Calabar, Southeastern Nigeria. Int J Virol 8:258–270

Lagenaria siceraria (Bottle gourd) Family: Cucurbitaceae

Vegetable

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

(CGMMV)

Lagenaria siceraria (Bottle gourd)

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CGMMV-infected plants of Lagenaria siceraria were reported from Saudi Arabia, Pakistan, India, China, and Myanmar (Capoor and Varma 1948; Vasudeva et al. 1949; Rao and Varma 1984; Al-Shahwan and Abdalla 1992; Zhang et al. 2009; Kim et al. 2010a; Ali et al. 2014; Amer 2015; Kumar et al. 2017; Nagendran et al. 2017). The virus-infected bottle gourd plants exhibit leaf mosaic and mottle symptoms. There is no known vector for this virus. The virus is transmissible by mechanical sap-inoculation, and by contact between plants. Transmission through contaminated irrigation water or nutrient solutions has also been reported. The virus is seed-transmitted in bottle gourd (Kim and Lee 2000). The virus is also pollen transmitted. For more details of CGMMV, refer to Cucumis sativus.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV was reported to infect plants of Lagenaria siceraria in India, Japan, and Serbia (Bhargava and Bhargava 1977; Al-Shahwan and Abdalla 1992; Takeshita et al. 2001; Dukic et al. 2006; Vucurovic et al. 2011; Nagendran et al. 2017). The affected bottle gourd plants exhibit greenish mosaic, veinbanding, blistering, yellowing, and leaf deformation symptoms on leaves. Plant height is reduced and the affected plants often produce distorted fruits. Different strains are known to cause different symptoms, and the movement protein and coat protein genes of CMV together determine the induction of severe chronic mosaic symptoms in bottle gourd (Takeshita et al. 2001). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(CABYV)

Family: Luteoviridae

CABYV infection in plants of Lagenaria siceraria was reported from China and Taiwan (Xiang et al. 2008; Knierim et al. 2010). The virus-infected bottle gourd plants exhibit yellowing of the older leaves, and leaves are thickened and brittle. The virus is transmitted by aphids, Myzus persicae, M. euphorbiae, and Aphis gossypii, in a persistent, circulative, non-propagative manner, (Lecoq et al. 1992). The virus is not transmissible by mechanical sap-inoculation. For more details of CABYV, refer to Cucumis melo.

Cucurbit chlorotic yellows virus

(CCYV)

Taxonomic position CCYV is a tentative member of the genus Crinivirus and family Closteroviridae CCYV infection in plants of Lagenaria siceraria was reported from Taiwan (Huang et al. 2010). The virus-infected bottle gourd plants exhibit symptoms of chlorosis, yellowing, and bleaching accompanied with green veins and brittleness on the lower leaves. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a semi-persistent manner. For more details of CCYV, refer to Cucumis melo.

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Lagenaria siceraria (Bottle gourd)

Grapevine fanleaf virus Taxonomic position Genus: Nepovirus

(GFLV)

Family: Secoviridae

GFLV infection in plants of Lagenaria siceraria cv. Turbinata was reported from Hungary (Horvath et al. 1994). The virus-infected bottle gourd plants exhibit symptoms of vein-clearing, yellow mosaic spots, and yellowing symptoms. The virus is transmitted by a nematode vector, Xiphinema index, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of GFLV, refer to Vitis vinifera.

Lagenaria mild mosaic virus Taxonomic position Genus: Potexvirus

(LaMMoV)

Family: Alphaflexiviridae

Geographical distribution LaMMoV infection in plants of Lagenaria siceraria was reported from Myanmar (Kim et al. 2010b). Symptoms and host(s) The virus-infected bottle gourd plants exhibit mosaic and mottle symptoms. Transmission The virus is mechanically sap-transmissible to a limited number of species in the Cucurbitaceae and Chenopodiaceae. No vector is known for this virus. Virion properties and genome The virions are filaments particles approximately 550 nm in length and 13 nm in diameter (Kim et al. 2010b). The genome consists of a single linear molecule of positive-sense ssRNA and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type, 18–27 kDa in size (Adams et al. 2004). A partial genomic sequence of 3860 nt is available (AB546335).

Lagenaria siceraria alphaendornavirus Taxonomic position Genus: Alphaendornavirus

(LsEV)

Family: Endornaviridae

Geographical distribution LsEV infection in plants of Lagenaria siceraria was reported from California and China (Kwon et al. 2014; Peng et al. 2017). Symptoms and host(s) The virus-infected bottle gourd plants do not exhibit any external symptoms.

Lagenaria siceraria (Bottle gourd)

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Transmission The virus is transmitted through the seed via both the ova and pollen. No horizontal spread has been observed in the field and no potential vectors have been identified. The virus is not mechanically sap-transmissible. Virion properties and genome The virus does not have a gene for a capsid protein; therefore no virion is formed. The genome is a double-stranded RNA of 15,088 bp (KF562072 = NC_023641) and contained one large ORF encoding a 576 kDa polyprotein (Kwon et al. 2014). The lengths of the 50 and 30 untranslated regions were 111 and 52 bp, respectively.

Papaya ringspot virus Taxonomic position Genus: Potyvirus

(PRSV)

Family: Potyviridae

PRSV infection in plants of Lagenaria siceraria was reported from India (Mantri et al. 2005). The virus-infected bottle gourd plants exhibit symptoms of mosaic, mottling, interveinal chlorotic bands, leaf distortion, malformation of fruits, and reduction in fruit size. In severe infections, blisters on the fruit surface commonly occurred. The virus is transmitted by an aphid vector Aphis gossypii in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and produces systemic mosaic mottling symptoms in cucurbitaceous and solanaceous hosts but failed to infect legumes. For more details of PRSV, refer to Carica papaya.

Soybean mosaic virus Taxonomic position Genus: Potyvirus

(SMV)

Family: Potyviridae

SMV infection in plants of Lagenaria siceraria was reported from India (Chakraborty et al. 2016). The virus-infected bottle gourd plants exhibit mosaic and yellowing symptoms on leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SMV, refer to Glycine max.

Squash mosaic virus Taxonomic position Genus: Comovirus

(SqMV)

Family: Secoviridae

SqMV infection in plants of Lagenaria siceraria was reported from Indonesia (Lestari and Nurhayati 2014). The virus is transmitted by beetle vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is seed-transmitted in Lagenaria siceraria at up to 100% (Lestari and Nurhayati 2014). For more details of SqMV, refer to Cucurbita pepo.

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Lagenaria siceraria (Bottle gourd)

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Lagenaria siceraria was reported from Thailand and India (Ito et al. 2008; Sohrab et al. 2010; Nagendran et al. 2016). The virus-infected bottle gourd plants exhibit symptoms of chlorotic mottling, mild curling, severe yellowing, and serious stunting. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is graft-transmissible. For more details of ToLCNDV, refer to Solanum lycopersicum.

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV has been reported to infect plants of Lagenaria siceraria in Serbia and India (Bhargava 1977; Chakraborty et al. 1997; Dukic et al. 2006). The virus-infected bottle gourd plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by also mechanical sap-inoculation. For more details of WMV, refer to Citrullus lanatus.

Zucchini yellow mosaic virus Taxonomic position Genus: Potyvirus

(ZYMV)

Family: Potyviridae

ZYMV infection in plants of Lagenaria siceraria was reported from India and Serbia (Verma et al. 2004; Dukic et al. 2006; Verma et al. 2006). The virus-affected bottle gourd plants showed severe mosaic, interveinal chlorosis, and leaf deformation that resulted in fern-leaf appearance and severe fruit distortion. The virus is transmitted by aphid vectors such as Aphis gossypii and Myzus persicae in a non-persistent manner, and also by mechanical sap-inoculation. The virus is seed-transmitted in Lagenaria siceraria at up to 13.3% (Lestari and Nurhayati 2014). For more details of ZYMV, refer to Cucurbita pepo.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Ali A, Hussain A, Ahmad M (2014) Occurrence and molecular characterization of Cucumber green mottle mosaic virus in cucurbit crops of KPK, Pakistan. Braz J Microbiol 45:1247–1253 Al-Shahwan IM, Abdalla OA (1992) A strain of Cucumber green mottle mosaic virus (CGMMV) from Bottlegourd in Saudi Arabia. J Phytopathol 134:152–156 Amer MA (2015) Biological and molecular characterization of Cucumber green mottle mosaic virus affecting bottle gourd and watermelon plants in Saudi Arabia. Int J Agric Biol 17:748–754 Bhargava B (1977) Some properties of two strains of Watermelon mosaic virus. Phytopathol Z 88:199–208 Bhargava B, Bhargava KS (1977) Cucurbit mosaic viruses in in Gorakhpur. Indian J Agri Sci 47:1–5

Lagenaria siceraria (Bottle gourd)

1343

Capoor SP, Varma PM (1948) Mosaic disease of Lagenaria vulgaris surveyed in the Bombay Province. Curr Sci 17:274–275 Chakraborty S, Sinha A, Pandey PK (1997) Occurrence of Watermelon mosaic virus – 1 on bottle gourd (Lagenaria siceraria Mol.) standl in Varanasi region. Veg Sci 24:147–149 Chakraborty P, Das S, Saha D, Saha A (2016) First report of Soybean mosaic virus infecting bottle gourd plant in India. Plant Dis 100:1509 Dukic N, Krstic B, Vico I, Berenji J, Duduk B (2006) First report of Zucchini yellow mosaic virus, Watermelon mosaic virus and Cucumber mosaic virus in bottlegourd (Lagenaria siceraria) in Serbia. Plant Dis 90:380 Horvath J, Tobias I, Hunyadi K (1994) New natural herbaceous hosts of grapevine fanleaf nepovirus. Hortic Sci 26: 31–32 Huang LH, Tseng HH, Li JT, Chen TC (2010) First report of Cucurbit chlorotic yellows virus infecting cucurbits in Taiwan. Plant Dis 94:1168 Ito T, Sharma P, Kittipakorn K, Ikegami M (2008) Complete nucleotide sequence of a new isolate of Tomato leaf curl New Delhi virus infecting cucumber, bottle gourd and muskmelon in Thailand. Arch Virol 153:611–613 Kim DH, Lee JM (2000) Seed treatment for Cucumber green mottle mosaic virus in gourd (Lagenaria siceraria) seeds and its detection. J Korean Soc Hortic Sci 41:1–6 Kim O-K, Mizutani T, Natsuaki KT, Lee K-W, Soe K (2010a) First report and the genetic variability of Cucumber green mottle mosaic virus occurring on Bottle Gourd in Myanmar. J Phytopathol 158:572–575 Kim O-K, Mizutani T, Soe K, Lee K-W, Natsuaki KT (2010b) Characterization of Lagenaria mild mosaic virus, a New Potexvirus from Bottle Gourd in Myanmar. Plant Dis 94:1225–1230 Knierim D, Deng TC, Tsai WS, Green SK, Kenyon L (2010) Molecular identification of three distinct Polerovirus species and a recombinant Cucurbit aphid-borne yellows virus strain infecting cucurbit crops in Taiwan. Plant Pathol 59:991–1002 Kumar A, Jailani AAK, Roy A, Mandal B (2017) The occurrence, biology and genomic properties of tobamoviruses infecting crop plants in India. In: Mandal B, Rao GP, Baranwal VK, Jain RK (eds) A century of plant virology in India. Springer Nature, Singapore. pp 429–443. ISBN 978-981-10-5671-0 Kwon SJ, Tan SH, Vidalakis G (2014) Complete nucleotide sequence and genome organization of an endornavirus from bottle gourd (Lagenaria siceraria) in California, USA. Virus Genes 49(1):163–168 Lecoq H, Bourdin D, Wipf-Scheibel C, Bon M, Lot H, Lemaire O, Herrbach E (1992) A new yellowing disease of cucurbits caused by a Luteovirus, Cucurbit aphid-borne yellows virus. Plant Pathol 41:749–761 Lestari SM, Nurhayati E (2014) Seed transmission efficiency of Squash mosaic virus on Cucurbitaceae. Fitopatol Indones 10:81–83 Mantri NL, Kitkaru AS, Misal MB, Ravi KS (2005) First report of Papaya ringspot virus-W in bottle gourd (Lagenaria siceraria) from India. Plant Pathol 54:806 Nagendran K, Karthika C, Parthasarathy S, Mohan Kumar S, Karthikeyan G (2016) Molecular evidence for the occurrence of Tomato leaf curl New Delhi virus on bottle gourd in Tamil Nadu, India. Veg Sci 43:73–77 Nagendran K, Mohankumar S, Aravintharaj R, Balaji CG, Manoranjitham SK, Singh AK, Rai AB, Singh B, Karthikeyan G (2017) The occurrence and distribution of major viruses infecting cucurbits in Tamil Nadu state, India. Crop Prot 99:10–16 Peng X, Pan H, Muhammad A, An H, Fang S, Li W, Zhang S (2017) Complete genome sequence of a new strain of Lagenaria siceraria endornavirus from China. Arch Virol 163:805–808 Rao DC, Varma A (1984) A strain of cucumber green mottle mosaic virus (CGMMV) from bottlegourd in Saudi Arabia. Phytopathol Z 109:325 Sohrab SS, Mandal B, Ali A, Varma A (2010) Chlorotic curly stunt: a severe begomovirus disease of bottle gourd in Northern India. Indian J Virol 21:56–63 Takeshita M, Suzuki M, Takanami Y (2001) Combination of amino acids in the 3a protein and the coat protein of cucumber mosaic virus determines symptom expression and viral spread in bottle gourd. Arch Virol 146: 697–711 Vasudeva RS, Raychaudhuri SP, Singh J (1949) A new strain of Cucumis virus 2. Indian Phytopathol 2:180–185 Verma R, Ahlawat YS, Tomer SPS, Prakash S, Pant RP (2004) First report of Zucchini yellow mosaic virus in bottle gourd (Lagenaria siceraria) in India. Plant Dis 88:426 Verma R, Baranwal VK, Prakash S, Tomer SPS, Singh J, Pant RP, Ahlawat YS (2006) Biological and molecular characterization of Zucchini yellow mosaic virus from naturally infected bottle gourd. Indian J Virol 17: 96–101 Vucurovic A, Bulajic A, Stankovic I, Ristic D, Berenji J, Krstic B (2011) Characterization of Cucumber mosaic virus originating from cucurbits in Serbia. Pestic Phytomed 26(4):325–336 Xiang HY, Shang QX, Han CG, Li DW, Yu JL (2008) First report on the occurrence of Cucurbit aphid-borne yellows virus on nine cucurbitaceous species in China. Plant Pathol 57:390 Zhang YJ, Li GF, Li MF (2009) Occurrence of Cucumber green mottle mosaic virus on cucurbitaceous plants in China. Plant Dis 93:200

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Lamium album (White dead-nettle)

Lamium album (White dead-nettle) Family: Lamiaceae

Leafy vegetable

Lamium mild mosaic virus Taxonomic position Genus: Fabavirus

(LMMV)

Family: Secoviridae

Geographical distribution LMMV infection in plants of Lamium album was first found in 1953 in the UK (Lovisolo 1958). The virus spreads in the UK and Spain (Lisa et al. 1982; Rangel et al. 2013). Symptoms and host(s) The virus-infected white dead-nettle plants exhibit mild mosaic symptoms. Transmission The virus is transmitted by aphids in a non-persistent manner. The virus is also mechanically saptransmissible. Virion properties and genome The virions are isometric, non-enveloped of two types, but similar in size 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is a bipartite, linear positive-sense, single-stranded RNA. The genome was the largest of the genus but maintained the typical organization, with RNA 1 of 6080 nt (KC595304 = NC_023016), RNA 2 of 4065 nt (KC595305 = NC_023017), and an unusually long 30 untranslated region in RNA 2 of 603 nt (Ikegami and Sharma 2011; Ferrer et al. 2007; Rangel et al. 2013; Sanfacon 2015; Thompson et al. 2017).

References Ferrer RM, Luis-Arteaga M, Guerri J, Moreno P, Rubio L (2007) Detection and identification of species of the genus Fabavirus by RT–PCR with a single pair of primers. J Virol Methods 144:156–160 Ikegami M, Sharma P (2011) Fabavirus. Comoviridae. In: Tidona C, Darai G (eds) The Springer Index of Viruses. Springer, New York, NY. https://doi.org/10.1007/978-0-387-95919-1 Lisa V, Luisoni E, Boccardo G, Milne R, Lovisolo O (1982) Lamium mild mosaic virus: a virus distantly related to broad bean wilt. Ann Appl Biol 100:467–476 Lovisolo O (1958) Virus e piante spontanee. Il Mosaico lieve del lamium nuovo virus di tipo maculatura anulare. Boll Staz Pat Veg Roma III Ser 15:89–137 Rangel EA, Ferriol I, Panno S, Davino S, Olmos A, Rubio L (2013) The complete genome sequence of Lamium mild mosaic virus, a member of the genus Fabavirus. Arch Virol 158(11):2405–2408 Sanfacon H (2015) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. In: Encyclopedia of Life Sciences. Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000764.pub3. http:// www.els.net Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J Gen Virol 98:529–531

Lamium maculatum (Spotted dead-nettle)

1345

Lamium amplexicaule (Henbit dead-nettle) Family: Lamiaceae

Leafy vegetable

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(CABYV)

Family: Luteoviridae

CABYV infection in plants of Lamium amplexicaule was reported from Tunisia (Mnari-Hattab et al. 2009). The virus-infected Henbit dead-nettle plants exhibit severe yellowing symptoms on older leaves. The virus is transmitted by aphid vectors, Myzus persicae, M. euphorbiae, and Aphis gossypii, in a circulative, non-propagative manner, and is not transmissible by mechanical sap-inoculation. For more details of CABYV, refer to Cucumis melo.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Lamium amplexicaule was reported from Korea (Kil et al. 2014). The virus-infected Henbit dead-nettle plants exhibit yellowing, stunting, and leaf curling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

References Kil EJ, Park J, Lee H, Kim J, Choi HS, Lee KY, Kim CS, Lee S (2014) Lamium amplexicaule (Lamiaceae): a weed reservoir for tomato yellow leaf curl virus (TYLCV) in Korea. Arch Virol 159(6):1305–1311 Mnari-Hattab M, Gauthier N, Zouba A (2009) Biological and molecular characterization of the Cucurbit aphid-borne yellows virus affecting cucurbits in Tunisia. Plant Dis 93:1065–1072

Lamium maculatum (Spotted dead-nettle) Family: Lamiaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Weed host

(CMV)

Family: Bromoviridae

CMV infection in plants of Lamium maculatum was reported from Bosnia and Herzegovina (Besta-Gajevic et al. 2013). The virus-infected spotted dead-nettle plants exhibit symptoms of severe mosaic accompanied

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Lathyrus cicera (Dwarf chickling)

by reddish brown necrosis and leaf deformation. The virus is transmitted by aphid vectors in a non-persistent manner and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Lamium leaf distortion virus Taxonomic position Genus: Caulimovirus

(LLDV)

Family: Caulimoviridae

Geographical distribution LLDV infection in plants of Lamium maculatum was reported from the USA (Zhang et al. 2008). Symptoms and host(s) The virus-infected spotted dead-nettle plants exhibit leaf distortion symptoms. Lamium maculatum is the only known host of this virus. Transmission The virus is not transmitted by mechanical or graft inoculation or by Myzus persicae. Virion properties and genome The virions are isometric, non-enveloped, and 45–50 nm in diameter with no obvious surface structure. The genome is a monopartite and circular, with dsDNA of 7713 bp, and consists of six open reading frames (EU554423 = NC_010737) (Zhang et al. 2008; Hohn 2011).

References Besta-Gajevic R, Jerkovic-Mujkic A, Pilic S, Stankovic I, Vucurovic A, Bulajic A, Krstic B (2013) Lamium maculatum is a natural host for Cucumber mosaic virus. Plant Dis 97:150 Hohn T (2011) Caulimovirus. Caulimoviridae. In: The Springer Index of Viruses. Springer, New York, pp 271–277. https://doi.org/10.1007/978-0-387-95919-1_41 Zhang L, Lockhart B, Dahal G, Olszewski N (2008) Studies on biology and genomic characterization of a caulimo-like virus associated with a leaf distortion disease of Lamium maculatum. Arch Virol 153:1181–1184

Lathyrus cicera (Dwarf chickling) Family: Fabaceae

Forage crop

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Lathyrus cicera was reported from Western Australia (Latham and Jones 2001). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Lathyrus odoratus (Sweet pea)

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Cucumber mosaic virus

(CMV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV infection in plants of Lathyrus cicera was reported from Western Australia (Latham and Jones 2001). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of CMV, refer to Cucumis sativus.

References Latham LJ, Jones RAC (2001) Incidence of virus infection in experimental plots, commercial crops and seed stocks of cool season crop legumes. Aust J Agric Res 52:397–413

Lathyrus clymenum (Spanish vetchling) Family: Fabaceae

Pulse crop

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Lathyrus clymenum was reported from Western Australia (Latham and Jones 2001). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

References Latham LJ, Jones RAC (2001) Incidence of virus infection in experimental plots, commercial crops and seed stocks of cool season crop legumes. Aust J Agric Res 52:397–413

Lathyrus odoratus (Sweet pea) Family: Fabaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Ornamental

(AMV)

Family: Bromoviridae

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Lathyrus odoratus (Sweet pea)

AMV infection in plants of Lathyrus odoratus was reported from the United Kingdom (Hull 1965). The virus-infected sweet pea plants exhibit symptoms of rosetting and twisting of the shoot apex, with chlorosis primarily at the margins of young leaves, dark necrotic streaks on the stem, and shortened, twisted peduncles. Streaking of the stem spreads rapidly, often leading to plant death. The virus is transmitted by a large number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Bean leafroll virus

(BLRV)

Synonyms Pea leaf roll virus Taxonomic position Genus: Luteovirus

Family: Luteoviridae

BLRV infection in plants of Lathyrus odoratus was reported from the United Kingdom (Hull 1965). The virus-infected sweet pea plants exhibit symptoms of rosetting of the apex, with upward rolling, crinkling, and thickening of the young leaves. Middle leaves may show interveinal chlorotic streaks. The virus is transmitted in a circulative, persistent manner, by aphids. The virus is not mechanically sap-transmissible. For more details of BLRV, refer to Phaseolus vulgaris.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Lathyrus odoratus was reported from the United Kingdom (Hull 1965). In sweet pea, the first symptoms are vein-clearing of young leaves, followed by interveinal mottle and then dark green vein-banding; interveinal chlorotic dots may also be seen, yielding a three-colored mottle. Light green streaks which may later become brown, and necrosis may spread up the stems, along petioles, and into leaf veins. Lighter-colored flower break may appear at the same time as early leaf symptoms, followed by dark streaks in later-formed flowers of pale-colored varieties. No flower break is seen in white-flowered varieties. The virus is transmitted non-persistently by aphids, but not transmitted through seeds of sweet pea. The virus is mechanically sap-transmissible. For more details of BYMV, refer to Phaseolus vulgaris.

Clover yellow vein virus Taxonomic position Genus: Potyvirus

(ClYVV)

Family: Potyviridae

ClYVV infection in plants of Lathyrus odoratus was reported from New Zealand (Forster and Musgrave 1985). The virus-infected sweet pea plants show leaf and stem necrosis and severe flower break. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ClYVV, refer to Trifolium spp.

Lathyrus odoratus (Sweet pea)

1349

Pea enation mosaic virus

(PEMV)

Taxonomic position PEMV is a tentative member of the genus Enamovirus and family Luteoviridae PEMV infection in plants of Lathyrus odoratus was reported from the United States, the United Kingdom, and Canada (Harju et al. 2011). The virus-infected sweet pea plants exhibit symptoms of mosaic and hyaline lesions with enations. The virus is transmitted by aphid vectors, Acyrthosiphon pisum and Myzus persicae, in the persistent manner, (Hull and Selman 1965; Getz et al. 1982). The virus is also mechanically transmissible and probably transmissible through seed. For more details of PEMV, refer to Pisum sativum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV-infected plants of Lathyrus odoratus were reported from Manitoba (Canada) and Hawaii (Cho et al. 1987). The virus-infected sweet pea plants exhibit symptoms of reddish brown to purple streaks which may extend to the full length of the stem. Circular to oval spots with diffuse margins are followed by yellowing and death of the leaves and stems. Blossoms sometimes develop circular patterns in the pigment. The virus is transmitted by a western flower thrips, Frankliniella occidentalis, in a persistentpropagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts and by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

White clover mosaic virus Taxonomic position Genus: Potexvirus

(WClMV)

Family: Alphaflexiviridae

WClMV infection in plants of Lathyrus odoratus was reported from the United Kingdom and New Zealand (Fry 1959; Hull 1965; Brown and Brunt 1988; Harju et al. 2011). The virus-infected sweet pea plants exhibit symptoms of chlorosis and mottle of young leaves, with severe necrotic streaking on the stem, leading to plant death within a few weeks (Brown and Brunt 1988). The virus is sap-transmissible and also transmissible through contact between plants. For more details of WClMV, refer to Trifolium spp.

References Brown JD, Brunt AA (1988) The natural occurrence of white clover mosaic virus in sweet pea (Lathyrus odoratus) in Southeast England. Plant Pathol 37:445–446 Cho JJ, Mau RFL, Mitchell WC, Gonsalves D, Yudin LS (1987) Host list of plants susceptible to Tomato spotted wilt virus (TSWV). Research extension series 078. Hawaii Institute of Tropical Agriculture and Human Resources, Honolulu Forster RLS, Musgrave BR (1985) Clover yellow vein virus in white clover (Trifolium repens) and sweet pea (Lathyrus odoratus) in the North Island of New Zealand. N Z Agric Res 28:575–578 Fry PR (1959) A clover mosaic virus in New Zealand pastures. N Z J Agric Res 2:971–981 Getz WM, Sylvester ES, Richardson J (1982) Pea enation mosaic virus transmission by the pea aphid: a multiphase model of virus transmission. Phytopathology 72:1145–1148

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1350

Lathyrus sativus (Grass pea)

Harju VA, Skelton AL, Monger WA, Forde SMD, Daly M, Nixon T, Lawson R, Bennett S, Mumford RA (2011) A report of viruses, viroids and phytoplasmas found in ornamental plants from 1999-2007: findings from Central Science Laboratory, UK. Acta Hortic 901:223–229 Hull R (1965) Virus diseases of sweet peas in England. Plant Pathol 14:150–153 Hull R, Selman IW (1965) The incidence and spread of viruses in sweet peas (Lathyrus odoratus L.) in relation to variety and the use of systemic insecticides. Ann Appl Biol 55:39–50

Lathyrus sativus (Grass pea) Family: Fabaceae

Pulse crop

Alfalfa mosaic virus

(AMV)

Taxonomic position Genus: Alfamovirus

Family: Bromoviridae

AMV infection in plants of Lathyrus sativus was reported from Western Australia (Latham and Jones 2001). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Lathyrus sativus was reported from Western Australia (Latham and Jones 2001). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Lathyrus sativus was reported from Western Australia (Latham and Jones 2001). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Pea enation mosaic virus

(PEMV)

Taxonomic position PEMV is a tentative member of the genus Enamovirus and family Luteoviridae

Lavandula spp.

1351

PEMV infection in plants of Lathyrus sativus was reported from Syria (Makkouk et al. 2001). The virus-infected grass pea plants exhibit symptoms of stunting, leaf mottling, and yellowing. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is mechanically sap-transmissible. For more details of PEMV, refer to Pisum sativum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Lathyrus sativus was reported from Manitoba and Canada (Zimmer and Haber 1992; Zimmer et al. 1992). The virus-infected grass pea plants exhibit symptoms of purplish discoloration of the pod, and straw-colored areas on stems, extending from the node up and down the stem. Severe infection resulted in complete ovule abortion, especially at the top node. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Latham LJ, Jones RAC (2001) Incidence of virus infection in experimental plots, commercial crops and seed stocks of cool season crop legumes. Aust J Agric Res 52:397–413 Makkouk KM, Kumari SG, Lesemann D-E (2001) First record of pea enation mosaic virus naturally infecting chickpea and grasspea crops in Syria. Plant Dis 85:1032 Zimmer RC, Haber S (1992) First report of tomato spotted wilt virus in Manitoba and of Lathyrus sativus as a host. Plant Dis 76:753 Zimmer RC, Myers K, Haber S, Campbell CG, Gubbels (1992) Tomato spotted wilt virus, a problem on grass pea and field pea in the greenhouse in 1990 and 1991. Can Plant Dis 72:29–31

Lavandula spp. Family: Lamiaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Medicinal

(AMV)

Family: Bromoviridae

AMV infection in plants of Lavandula spp. was reported from Croatia, France, Spain, New Zealand, and Italy (Marchoux and Rougier 1974; Fletcher 1987; Martinez-Priego et al. 2004; Bruni et al. 2006; Parrella et al. 2010; Vrandecic et al. 2013; Stankovic et al. 2014; Mitrofanova et al. 2018). The virusinfected lavandula plants exhibit bright yellow calico mosaic, leaf distortion, and growth reduction symptoms. The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

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1352

Leersia oryzoides (Cutgrass)

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Lavandula angustifolia was reported from Poland and Northern Italy (EU432182) (Kobylko et al. 2008; Davino et al. 2012; Mitrofanova et al. 2018). The virus-infected lavandula plants exhibited yellow mottling, leaf deformation, and growth reduction. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

References Bruni R, Bellardi MG, Parrella G, Bianchi A (2006) Impact of alfalfa mosaic virus subgroup I and II isolates on terpene secondary metabolism of Lavandula vera D.C., Lavandulaalardii and eight cultivars of L. hybrida. Rev Physiol Mol Plant Pathol 68:189–197 Davino S, Panno S, Rangel EA, Davino M, Bellardi MG, Rubio L (2012) Population genetics of Cucumber mosaic virus infecting medicinal, aromatic and ornamental plants from northern Italy. Arch Virol 157:739–745 Fletcher JD (1987) New plant disease records in New Zealand: additional hosts of alfalfa mosaic virus and cucumber mosaic virus. N Z J Agric Res 30:505–506 Kobylko T, Danda P, Hasiow B, Borodynko N, Pospieszny H (2008) First report of Cucumber mosaic virus on Lavandula angustifolia in Poland. Plant Dis 92:978 Marchoux G, Rougier J (1974) Alfalfa mosaic virus: isolation from lavandin (Lavandula hybrida Rev.) and eggplant (Solanum melongena L.). Ann Phytopathol 6:191–196 Martinez-Priego L, Córdoba MC, Jordá C (2004) First report of Alfalfa mosaic virus in Lavandula officinalis. Plant Dis 88(8):908 Mitrofanova IV, Zakubanskiy AV, Mitrofanova OV (2018) Viruses infecting main ornamental plants: an overview. Ornamental Hortic 24:95–102 Parrella G, Acanfora N, Bellardi MG (2010) First record and complete nucleotide sequence of Alfalfa mosaic virus from Lavandula stoechas in Italy. Plant Dis 94(7):924 Stankovic I, Vrandecic K, Cosic J, Milojevic K, Bulajic A, Krstic B (2014) The spreading of Alfalfa mosaic virus in lavandin in Croatia. Pestic Phytomed 29:115–122 Vrandecic K, Jurkovic D, Cosic J, Stankovic I, Vucurovic A, Bulajic A, Krstic B (2013) First report of Alfalfa mosaic virus infecting Lavandula x intermedia in Croatia. Plant Dis 97:1002

Leersia oryzoides (Cutgrass) Family: Poaceae

Weed host

Barley yellow dwarf virus

(BYDV)

Taxonomic position BYDV is a tentative member of the genus Luteovirus and family Luteoviridae BYDV infection in plants of Leersia oryzoides was reported from Northern Italy (Osler et al. 1980). The virus is transmitted by aphid vectors, Rhopalosiphum padi and Sitobion avenae, in a circulative, nonpropagative manner, suggesting the probability that it was BYDV-PAV. No mechanical transmission of this virus is reported. For more details of BYDV, refer to Hordeum vulgare.

Lens culinaris (Lentil)

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References Osler R, Amici A, Longoni CE (1980) Leersia oryzoides, a natural host and winter reservoir of the rice “Giallume” strain of barley yellow dwarf virus. J Phytopathol 97:242–251

Lens culinaris (Lentil) Family: Fabaceae

Pulse crop

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

(AMV)

Family: Bromoviridae

AMV infection occurs wherever Lens culinaris crops are grown (Kaiser 1981; Bos et al. 1988; Jones and Coutts 1996; Lathman and Jones 2001; Makkouk et al. 2003a; Kumari et al. 2009b; Chatzivassiliou et al. 2016). The virus-infected lentil plants exhibit mild chlorosis with leaves reduced in size and frequently malformed. Necrosis often occurs along leaflet margins. Plants are usually stunted and may die prematurely. Flowering and pod formation are reduced and seeds are small and shriveled. The virus is transmitted by a large number of aphid species in a non-persistent manner. The virus is mechanically sap-transmissible. The virus is seed-transmitted in Lens culinaris which was up to 0.1–5% (Jones and Coutts 1996; Makkouk and Attar 2003). For more details of AMV, refer to Medicago sativa.

Bean leafroll virus Taxonomic position Genus: Luteovirus

(BLRV)

Family: Luteoviridae

Lens culinaris plants infected naturally with BLRV were first observed in Iran by Kaiser et al. (1968). The virus spreads in Australia, Europe, Africa, Bangladesh, Ethiopia, Iran, Iraq, Syria, Tunisia, Greece, and the USA (Kaiser 1973; Bos et al. 1988; Klein et al. 1991; Makkouk et al. 1992, 2003a; Bakr 1993; Tadesse et al. 1999a; El-Muadhidi et al. 2001; Kumari 2002; Mustafayev et al. 2011; Chatzivassiliou et al. 2016). The virus infection in lentil produces interveinal chlorosis, yellowing, stunting, leaf rolling, reddening and thickening of the leaves, and suppression of flowering and pod set. Entire leaflets can turn red in cool temperatures. The virus is transmitted only by aphid vectors, Acyrthosiphon pisum, Aphis craccivora, Aphis fabae, and Myzus persicae, in a persistent, non-propagative manner, (Kaiser 1973). The virus is not mechanically sap-transmissible, and there are reports of transmission through seed and pollen. For more details of BLRV, refer to Phaseolus vulgaris.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

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Lens culinaris (Lentil)

BYMV was first reported in plants of Lens culinaris by Kaiser et al. (1968) from Iran. The virus occurs on lentil in Bangladesh, Egypt, Ethiopia, Iran, Iraq, Jordan, New Zealand, Syria, the USA, Puerto Rico, Greece, and Turkey (Kaiser 1973, 1981; Russo et al. 1981; Bos et al. 1988; Makkouk et al. 1992, 1993, 2003a; Bakr 1993; Fletcher 1993; Bayaa et al. 1998; Tadesse et al. 1999a; Al-Mabrouk and Mansour 2000; El-Muadhidi et al. 2001; Chatzivassiliou et al. 2016). The virus-infected lentil plants exhibit chlorosis, mild mosaic or mottling, and stunting. Leaves often become twisted or curled with necrosis along the margins. Flowering and pod formation are reduced as a result of infection, and consequently little seed is produced. The virus is transmitted by aphid vectors, Acyrthosiphon pisum, Aphis fabae, A. craccivora, Myzus persicae, and Macrosiphum euphorbiae, in a non-persistent manner (Edwardson and Christie 1986). The virus is also mechanically sap-transmissible to a large number of hosts. Seed-transmission is also recorded up to 0.8% in lentil (Kumari et al. 1994; Jones and Coutts 1996). For more details of BYMV, refer to Phaseolus vulgaris.

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

BWYV occurs in plants of Lens culinaris in Ethiopia, Iran, New Zealand, Pakistan, Syria, and Turkey (Fletcher 1993; Tadesse et al. 1999a, b; Makkouk et al. 1992, 1998, 2001, 2002, 2003a; Kumari 2002; Kumari et al. 2009b; Mustafayev et al. 2011). The virus infection on lentil induces symptoms such as yellowing or bronzing with leaflets becoming leathery and brittle. Usually, leaf margins become reddish, but the entire leaflets may turn red under cool conditions. Plants are severely stunted or may die if infected early. Internodes become shortened in the upper parts of plants infected at a later growth stage. Pods in the infected plants are reduced in size and poorly filled or may remain seedless. The virus is transmitted by aphid vectors, Myzus persicae, Aphis craccivora, Acyrthosiphon pisum, and Aulacorthum solani, in a circulative, non-propagative manner, (Boswell and Gibbs 1983). The virus is not transmissible by mechanical sap-inoculation. For more details of BWYV, refer to Beta vulgaris.

Broad bean stain virus Taxonomic position Genus: Comovirus

(BBSV)

Family: Secoviridae

BBSV infection in plants of Lens culinaris was reported from North America, West Asia, Ethiopia, Iran, Jordan, Syria and Turkey, Australia, and New Zealand (Makkouk et al. 1987, 1992, 2003a; Bos et al. 1988; Kumari et al. 1993; Bayaa et al. 1998; Tadesse et al. 1999a; Al-Mabrouk and Mansour 2000). The virus-infected lentil plants show very mild mottling symptoms. BBSV is transmitted primarily by beetle species Apion aestivum, A. arrogans, Sitona crinita, S. limosa, and S. lineatus in a non-persistent manner (Cockbain et al. 1975; Makkouk and Kumari 1989, 1995). The virus is mechanically saptransmissible and seed-transmitted up to 2.1% (Makkouk and Azzam 1986). For more details of BBSV, refer to Vicia faba.

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Chickpea chlorotic dwarf virus Taxonomic position Genus: Mastrevirus

(CpCDV)

Family: Geminiviridae

CpCDV occurs in plants of Lens culinaris in India, Syria, Pakistan, and Iran (Makkouk et al. 2001, 2002, 2003a; Kumari et al. 2009b). The virus-infected lentil plants show reddening or yellowing of leaflets with severe stunting. The virus is reported to naturally infect chickpea, lentil, faba bean, phaseolus bean, and sugar beet. The virus is transmitted by leafhopper vectors, Orosius orientalis and Orosius albicinctus, in a persistent, circulative, and non-propagative manner. The virus is not mechanically sap-transmissible. For more details of CpCDV, refer to Cicer arietinum.

Chickpea chlorotic stunt virus Taxonomic position Genus: Polerovirus

(CpCSV)

Family: Luteoviridae

CpCSV infection in plants of Lens culinaris was reported from Azerbaijan and Syria (Asaad et al. 2009; Kumari et al. 2009b; Mustafayev et al. 2011). The virus-infected lentil plants exhibit yellowing, reddening, and/or stunting symptoms. The virus is transmitted by aphid vectors in a circulative, nonpropagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of CpCSV, refer to Cicer arietinum.

Clover yellow vein virus Taxonomic position Genus: Potyvirus

(ClYVV)

Family: Potyviridae

ClYVV infection in plants of Lens culinaris was reported from Spain (Ortiz et al. 2009). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ClYVV, refer to Trifolium spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV was first recorded to infect plants of Lens culinaris in Iran (Kaiser et al. 1968). The virus naturally infects lentil in Australia, Ethiopia, Greece, India, Iran, Nepal, New Zealand, Pakistan, and Syria (Kaiser 1973; Rangaraju and Chenulu 1981; Fletcher 1993; Karki 1993; Bashir et al. 1994; Mouhanna et al. 1994; Jones and Coutts 1996; Tadesse et al. 1999a; Lathman and Jones 2001; Makkouk et al. 2001, 2003b; Chatzivassiliou et al. 2016). CMV-infected lentil plants exhibit symptoms of general chlorosis of the plant, leaf malformation, and stunting, often followed by premature death. Leaflets may show a mild mosaic or reddening of leaf margins. A reduction in pod number or filling of pods generally occurs resulting in yield loss. The virus is transmitted by aphid vectors in a non-persistent manner, and

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Lens culinaris (Lentil)

also by mechanical sap-inoculation. The virus is seed-transmitted up to 64.2% in Lens culinaris (Jones and Coutts 1996; Fletcher et al. 1999; Makkouk and Attar 2003; Makkouk et al. 2003b). For more details of CMV, refer to Cucumis sativus.

Faba bean necrotic yellows virus Taxonomic position Genus: Nanovirus

(FBNYV)

Family: Nanoviridae

FBNYV occurs in plants of Lens culinaris in Azerbaijan, Egypt, Ethiopia, Iran, Iraq, Pakistan, Spain, Syria, and Turkey (Makkouk et al. 1992, 1998, 2001, 2002, 2003a; Bayaa et al. 1998; Tadesse et al. 1999a; El-Muadhidi et al. 2001; Kumari et al. 2009a; Mustafayev et al. 2011). The virus-infected lentil plants exhibit leaves becoming thick and brittle and show interveinal chlorotic blotches starting from the leaf margins. The uppermost young leaves remain very small and cupped upward, whereas the older leaves are rolled downward. New shoots, leaves, and flowers develop poorly. Approximately after 3–4 weeks, infection is established, interveinal chlorosis becomes necrotic, and plants then die within 5–7 weeks. The virus is transmitted by aphid vectors, such as Acyrthosiphon pisum, Aphis craccivora, and A. fabae, in a persistent but non-propagative manner (Franz et al. 1998; Makkouk et al. 1998). The virus is not mechanically sap-transmissible and has negative results were obtained for pollen and seedtransmission. For more details of FBNYV, refer to Vicia faba.

Lentil stunt virus Taxonomic position The virus is a tentative member of the family Luteoviridae. Geographical distribution The virus infection in plants of Lens culinaris was reported from Northeast Africa (Abraham et al. 2008). Symptoms and host(s) The virus-infected lentil plants exhibit stunting symptoms. Virion properties and genome The genome is a linear single-stranded RNA of ca. 6 kb that has five to six major open reading frames (ORFs); a partial sequence of 581 nt (GQ118152) is available (Abraham et al. 2008).

Pea enation mosaic virus 1 Taxonomic position Genus: Enamovirus

(PEMV-1)

Family: Luteoviridae

PEMV-1 infection in plants of Lens culinaris was reported from Syria, Iran, Greece, and the USA (Klein et al. 1991; Makkouk et al. 1999, 2003a; Kumari et al. 2001; Chatzivassiliou et al. 2016). The

Lens culinaris (Lentil)

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virus-infected lentil plants exhibit rolling of leaves accompanied by mottling and tip wilting or necrosis. The virus is transmitted by aphid vectors in a circulative, non-propagative manner (dependent upon coinfection with the luteovirus PEMV-2) and is also mechanically sap-transmissible (whereas PEMV-2 is not mechanically transmissible). For more details of PEMV-1 and PEMV-2, refer to Pisum sativum.

Pea seed-borne mosaic virus Taxonomic position Genus: Potyvirus

(PSbMV)

Family: Potyviridae

PSbMV infection in plants of Lens culinaris was reported from West Asia, North Africa, Syria, Western Australia, Greece, the UK, and the USA (Hampton 1982; Goodell and Hampton 1984; Alconero et al. 1986; Lathman and Jones 2001; Kumari et al. 1993; Makkouk et al. 1992, 1993, 2001, 2003a; Al-Mabrouk and Mansour 1998, 2000; Mustafayev et al. 2011; Van Leur et al. 2013; Giakountis et al. 2015; Chatzivassiliou et al. 2016). The virus-infected lentil plants show leaves which are narrowed, reduced in size with mosaic and mottling, and chlorosis or necrosis of shoot tips. A downward leaf roll is common and plants are usually stunted. Symptoms may vary in lentil due to cultivar differences, virus strain, and environmental effects. Seed pods may be abnormal in shape and size with reduced fewer seed. The virus is transmitted by aphid vectors, Aphis pisum, A. craccivora, and A. fabae, in a non-persistent manner. The virus is mechanically sap-transmissible and also transmitted through the seed up to 44% (Hampton and Muehlbauer 1977; Eppler et al. 1988; Varma et al. 1991; Kumari and Makkouk 1995; Bayaa et al. 1998; Makkouk et al. 2001; Coutts et al. 2008). For more details of PSbMV, refer to Pisum sativum.

Pea streak virus Taxonomic position Genus: Carlavirus

(PeSV)

Family: Betaflexiviridae

PeSV infection in plants of Lens culinaris was reported from the USA, Puerto Rico, North America, and Europe (Kaiser 1981; Bos et al. 1988; Kumari et al. 2009b). The virus infection in lentil results in wilting of the terminal shoots and vascular discoloration, a condition that contributes to plant death, especially at the seedling stage. Infected plants exhibit stunting, yellowing of leaflets, and wilting. Pods that do form usually are poorly filled, seed is misshapen, and yield losses can be significant. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation; no seed and pollen transmission is reported. For more details of PeSV, refer to Pisum sativum.

Potato leafroll virus Taxonomic position Genus: Polerovirus

(PLRV)

Family: Luteoviridae

PLRV infection in plants of Lens culinaris was reported from Syria (Makkouk et al. 1992). The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of PLRV, refer to Solanum tuberosum.

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Red clover vein mosaic virus Taxonomic position Genus: Carlavirus

(RCVMV)

Family: Betaflexiviridae

RCVMV infection in plants of Lens culinaris was reported from the Pacific Northwest of the USA (Larsen and Myers 1998; Kumari et al. 2009b). The virus-infected lentil plants exhibit symptoms of severe stunting, proliferation of axillary branches, and general chlorosis or death. The virus is transmitted by the pea aphid, Acyrthosiphon pisum, in a non-persistent manner, and also by mechanical sap-inoculation to a number of hosts. For more details of RCVMV, refer to Trifolium spp.

Soybean dwarf virus

(SbDV)

Synonyms Subterranean clover red leaf virus Taxonomic position Genus: Luteovirus

Family: Luteoviridae

SbDV infection in plants of Lens culinaris was reported from Australia, New Zealand, Iran, and Syria (Makkouk et al. 1997, 2002; Tadesse et al. 1999b; Kumari et al. 2009b). The virus-infected lentil plants exhibit yellowing and severe stunting symptoms. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by either mechanical inoculation or by contact between plants. For more details of SbDV, refer to Glycine max.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Lens culinaris was reported from Brazil (Fonseca et al. 1995; Kumari et al. 2009b). The virus-infected lentil plants exhibit symptoms of chlorosis and malformation of the apical leaves, ring-spot lesions on pods, and stunting. The virus is transmitted by thrips vectors in a persistentpropagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Abraham AD, Varrelmann M, Vetten HJ (2008) Molecular evidence for the occurrence of two new luteoviruses in cool season food legumes in northeastern Africa. Afr J Biotechnol 7:414–420 Alconero R, Provvidenti R, Gonsalves D (1986) Three Pea seed-borne mosaic pathotypes from pea and lentil germplasm. Plant Dis 70:783–786 Al-Mabrouk O, Mansour AN (1998) Viruses affecting lentil in Jordan. Indian Phytopathol 51:1–9 Al-Mabrouk O, Mansour AN (2000) Viruses affecting lentil in Jordan. Arab J Plant Protect 18:103–104

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Asaad NY, Kumari SG, Haj-Kassem AA, Shalaby A-BA, Al-Shaabi S, Malhotra RS (2009) Detection and characterization of Chickpea Chlorotic Stunt Virus in Syria. J Phytopathol 157:756–761 Bakr MA (1993) Plant protection of lentil Bangladesh. In: Erskine W, Saxena MC (eds) Lentil in South Asia, Proceedings of the seminar on lentils in South Asia, 11–15 March 1991, New Delhi, India. International Center for Agricultrual Research in the Dry Areas (ICARDA), Aleppo, pp 177–184 Bashir M, Tahir M, Malik BA (1994) Natural occurrence of Cucumber mosaic virus in lentil in Pakistan. Lens Newsl 21(1):44–45 Bayaa B, Kumari SG, Akkaya A, Erskine W, Makkouk KM, Turk Z, Ozberk I (1998) Survey of major biotic stresses of lentil in South – East Anatolia, Turkey. Phytopathol Mediterr 37:88–95 Bos L, Hampton RO, Makkouk KM (1988) Viruses and virus diseases of pea, lentil, faba bean and chickpea. In: Summerfield RJ (ed) World crops: cool season food legumes. Kluwer Academic Publishers, Dordrecht, pp 591–615 pp 1179 Boswell KF, Gibbs J (1983) Viruses of legumes: descriptions and keys from virus identification and data exchange. Australian National University, Canberra, 139 pp Chatzivassiliou EK, Giakountis A, Kumari SG, Makkouk KM (2016) Viruses affecting lentil (Lens culinaris Medik.) in Greece; incidence and genetic variability of Bean leafroll virus and Pea enation mosaic virus. Phytopathol Mediterr 55:239–252 Cockbain AJ, Bowen R, Etheridge P (1975) Attempts to control the spread of BBSV/EAMB. Rep Rothamsted Exp Stn 1974:235–236 Coutts BA, Prince RT, Jones RAC (2008) Further studies on Pea seed-borne mosaic virus in cool-season crop legumes: responses to infection and seed quality defects. Aust J Agric Res 59:1130–1145 Edwardson JR, Christie RG (1986) Viruses infecting forage legumes. Monograph 14, 3 vols. Florida Agricultural Experiment Stations, Gainesville., 742 pp. El-Muadhidi MA, Makkouk KM, Kumari SG, Jerjess M, Murad SS, Mustafa RR, Tarik F (2001) Survey for legume and cereal viruses in Iraq. Phytopathol Mediterr 40:224–233 Eppler A, Kheder MA, Schlosser E (1988) Viruses in lentils raised from seeds collected on local markets in Egypt. In: Proceedings of the 5th international congress of plant pathology, Kyoto, Japan. (Abstr.) Fletcher JD (1993) Surveys of virus disease in pea, lentil, dwarf and broad bean crops in South Island, New Zealand. N Z J Crop Hortic Sci 21:45–53 Fletcher JD, Russell AC, Butler RC (1999) Seed-borne Cucumber mosaic virus in New Zealand lentil crops: yield effects and disease incidence. N Z J Crop Hortic Sci 27:197–204 Fonseca MEN, Doiteux LS, de Avila AC, Lima MI, Kitajima EW (1995) Detection of Tomato spotted wilt tospovirus in lentil. Plant Dis 79:320 Franz A, Makkouk KM, Vetten HJ (1998) Acquisition, retention and transmission of Faba bean necrotic yellows virus by two of its aphid vectors, Aphis craccivora (Koch) and Acyrthosiphon pisum (Harris). J Phytopathol 146(7):347–355 Giakountis A, Skoufa A, Paplomatas EI, Tokatlidis IS, Chatzivassiliou EK (2015) Molecular characterization and phylogenetic analysis of a Greek lentil isolate of Pea seedborne mosaic virus. Phytoparasitica 43:615–628 Goodell JJ, Hampton RO (1984) Ecological characteristics of the lentil strain of Pea seed-borne mosaic virus. Plant Dis 68:148–150 Hampton RO (1982) Incidence of the lentil strain of Pea seed-borne mosaic virus as a contaminant of Lens culinaris germplasm. Phytopathology 72:695–698 Hampton RO, Muehlbauer FJ (1977) Seed transmission of Pea seed-borne mosaic virus in lentils. Plant Dis Reptr 61:235–238 Jones RAC, Coutts BA (1996) Alfalfa mosaic and Cucumber mosaic virus infection in chickpea and lentil: incidence and seed transmission. Ann Appl Biol 129:491–506 Kaiser WJ (1973) Etiology and biology of viruses affecting lentil (Lens esculenta Moench.) in Iran. Phytopathol Mediterr 12:7–14 Kaiser WJ (1981) Disease of chickpea, lentil, pigeonpea and tepary bean in continental United States and Puerto Rico. Econ Bot 35:300–320 Kaiser WJ, Danesh D, Okhovat M, Mossahebi H (1968) Diseases of pulse crops (edible legumes) in Iran. Plant Dis Reptr 52:687–691 Karki PB (1993) Plant protection of lentil in Nepal. In: Erskine W, Saxena MC (eds) Lentil in South Asia, Proceedings of the seminar on lentils in South Asia, 11–15 March 1991, New Delhi, India. International Center for Agricultural Research in the Dry Areas (ICARDA), Aleppo, pp 187–191 Klein RE, Larsen RC, Kaiser WJ (1991) Virus epidemic of grain legume in eastern Washington. Plant Dis 75:1186 Kumari SG (2002) A study on luteoviruses affecting cool-season food legumes. PhD thesis, Aleppo University, Aleppo, Syria, 230 pp Kumari SG, Makkouk KM (1995) Variability among twenty lentil genotypes in seed transmission rated and yield loss induced by pea seed-borne mosaic potyvirus infection. Phytopathol Mediterr 34:129–132

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Kumari SG, Makkouk KM, Ismail ID (1993) Survey of seed-borne viruses in lentil in Syria and their effects on lentil yield. Arab J Plant Protect 11:28–32 Kumari S, Makkouk K, Ismail ID (1994) Seed transmission and yield loss induced in lentil by Bean yellow mosaic potyvirus. Lens Newsl 21:42–44 Kumari SG, Makkouk KM, Bayaa B (2001) Pea enation mosaic virus-1 infecting lentil in Syria, and further information on its host range, purification, serology and transmission characteristics. Arab J Plant Protect 19(2):65–72 Kumari SG, Attar N, Mustafayev E, Akparov Z (2009a) First report of Faba bean necrotic yellows virus affecting legume crops in Azerbaijan. Plant Dis 93:1220 Kumari SG, Larsen R, Makkouk KM, Bashir M (2009b) Virus diseases of lentil and their control. In: Erskine W, Muehlbauer F, Sarker A, Sharma B (eds) The lentil botany, production and uses. CAB International, p 325 Larsen RC, Myers JR (1998) First report of red clover vein mosaic carlavirus naturally infecting lentil. Plant Dis 82:1064 Latham LJ, Jones RAC (2001) Incidence of virus infection in experimental plots, commercial crops and seed stocks of cool season crop legumes. Aust J Agric Res 52:397–413 Makkouk KM, Attar N (2003) Seed transmission of Cucumber mosaic virus and Alfalfa mosaic virus in lentil seeds. Arab J Plant Prot 21:49–52 Makkouk KM, Azzam OI (1986) Detection of broad bean stain virus in lentil seed groups. LENS Newsl 13:37–38 Makkouk KM, Kumari SG (1989) Apion arrogans, a weevil vector of broad bean mottle virus. FABIS Newsl 25:26–27 Makkouk KM, Kumari SG (1995) Transmission of broad bean stain comovirus and broad bean mottle bromovirus by weevils in Syria. J Plant Dis Protect 102(2):136–139 Makkouk KM, Bos L, Azzam OI, Katul L, Rizkallah A (1987) Broad bean stain virus: identification, detectability with ELISA in faba bean leaves and seeds, occurrence in West Asia and North Africa, and possible wild hosts. Neth J Plant Pathol 93:97–106 Makkouk KM, Kumari SG, Al-Daoud E (1992) Survey of viruses affecting lentil (Lens culinaris Med.) in Syria. Phytopathol Mediterr 31:188–190 Makkouk KM, Kumari SG, Bos L (1993) Pea seed-borne mosaic virus: occurrence in faba bean (Vicia faba) and lentil (Lens culinaris) in West Asia and North Africa, and further information on host range, purification, serology and transmission characteristics. Neth J Plant Pathol 99:115–124 Makkouk KM, Damsteegt V, Johnstone GR, Katul L, Lesemann D-E, Kumari SG (1997) Identification and some properties of Soybean dwarf luteovirus affecting lentil in Syria. Phytopathol Mediterr 36:135–144 Makkouk KM, Bashir M, Jones R (1998) First record of Faba bean necrotic yellows virus and Beet western yellows luteovirus affecting lentil and chickpea in Pakistan. Plant Dis 82:591 Makkouk KM, Kumari SG, Bayaa B (1999) First report of Pea enation mosaic virus affecting lentil (Lens culinaris Medik.) in Syria. Plant Dis 83:303 Makkouk KM, Bashir M, Jones RAC, Kumari SG (2001) Survey for viruses in lentil and chickpea crops in Pakistan. Z Pflanzenkrankh Pflanzenschutz 108:258–268 Makkouk KM, Fazlali Y, Kumari SG, Farzadfar S (2002) First record of Beet western yellows virus, Chickpea chlorotic dwarf virus, Faba bean necrotic yellows virus and Soybean dwarf virus infecting chickpea and lentil crops in Iran. Plant Pathol 51:387 Makkouk KM, Kumari SG, Shahreen N, Fazlali Y, Farzadfar S, Ghotbi T, Reza Mansouri A (2003a) Identification and seasonal variation of viral diseases of chickpea and lentil in Iran. J Plant Dis Protect 110:157–169 Makkouk KM, Attar N, Kumari SG (2003b) Seed transmission of Cucumber mosaic virus in lentil seeds in Syria. In: Eighth Arab congress of plant protection, El-Beida, Libya. p 59 Mouhanna AM, Makkouk KM, Ismail ID (1994) Survey of virus disease of wild and cultivated legumes in the coastal region of Syria. Arab J Plant Protect 12(1):12–19 Mustafayev E, Kumari SG, Attar N, Zeynal A (2011) Viruses infecting chickpea and lentil crops in Azerbaijan. Australas Plant Pathol 40(6):612–620 Ortiz V, Castro S, Romero J (2009) First report of Clover yellow vein virus in grain legumes in Spain. Plant Dis 93:106 Rangaraju R, Chenulu VV (1981) Occurrence of interveinal chlorosis of lentil in India. Curr Sci 50:191–192 Russo M, Kishtah AA, Tolba MA (1981) A disease of lentil caused by Bean yellow mosaic virus in Egypt. Plant Dis 65:611–612 Tadesse N, Ali K, Gorfu D, Yusuf A, Abraham A, Ayalew M, Lencho A, Makkouk KM, Kumari SG (1999a) Survey for chickpea and lentil virus diseases in Ethiopia. Phytopathol Mediterr 38:149–158 Tadesse N, Ali K, Gorfu D, Abraham A, Lencho A, Ayalew M, Yusuf A, Makkouk KM, Kumari SG (1999b) First report of Soybean dwarf virus infecting lentil and Beet western yellows virus infecting lentil and chickpea crops in Ethiopia. Plant Dis 83:589 Van Leur JAG, Freeman AE, Aftab M, Spackman M, Redden B, Matern M (2013) Identification of seed-borne Pea seedborne mosaic virus in lentil (Lens culinaris) germplasm and strategies to avoid its introduction in commercial Australian lentil fields. Australas Plant Dis Notes 8:75–77 Varma A, Khetarpal RK, Viswanath SM, Dhirendra K, Maury Y, Sharma B, Tyagi MC, Kumar D (1991) Detection of Pea seed-borne mosaic virus in commercial seeds of pea and germplasm of pea and lentil. Indian Phytopathol 44:107–111

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Leonurus sibiricus (Honeyweed, Motherwort) Family: Lamiaceae

Medicinal

Beet western yellows virus Taxonomic position Genus: Polerovirus

(BWYV)

Family: Luteoviridae

BWYV infection in plants of Leonurus sibiricus was reported from Korea (Kwon et al. 2016). The virus-infected honeyweed plants exhibit mild mottle-mosaic and chlorosis symptoms. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical inoculation. For more details of BWYV, refer to Beta vulgaris.

Broad bean wilt virus 2 Taxonomic position Genus: Fabavirus

(BBWV-2)

Family: Secoviridae

BBWV-2 infection in plants of Leonurus sibiricus was reported from Korea (Seo et al. 2014). The virusinfected honeyweed plants exhibit mosaic, yellowing, and stunting symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BBWV-2, refer to Vicia faba.

Euphorbia yellow mosaic virus Taxonomic position Genus: Begomovirus

(EuYMV)

Family: Geminiviridae

EuYMV infection in plants of Leonurus sibiricus was reported from Brazil (Ferro et al. 2017). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of EuYMV, refer to Euphorbia heterophylla.

Leonurus mosaic virus Taxonomic position Genus: Begomovirus

(LeMV)

Family: Geminiviridae

Geographical distribution LeMV infection in plants of Leonurus sibiricus was reported from Brazil (Faria and Maxwell 1999; Fonseca et al., unpublished - JQ429791).

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Symptoms and host(s) The virus-infected honeyweed plants exhibit mosaic symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, measuring about 22  38 nm, with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2652 nt (JQ429791 = NC_038450) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Motherwort yellow mottle virus Taxonomic position Genus: Torradovirus

(MYMoV)

Family: Secoviridae

Geographical distribution MYMoV infection in plants of Leonurus sibiricus was reported from Korea (Seo et al. 2015; van der Vlugt et al. 2015). Symptoms and host(s) The virus-infected motherwort plants exhibit yellow mottle, mild mosaic, and stunting symptoms. Transmission The virus is transmitted by whitefly vectors in a semi-persistent manner, (Verbeek et al. 2014). The virus is also mechanically sap-transmissible. Virion properties and genome The virions are isometric, non-enveloped of two types, but similar in size 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome of this virus is composed of two single-stranded RNAs carrying poly (A) tails. RNA1 (7068 nt; KM22970 = NC_035218) contains one large open reading frame (RNA1-ORF1), while RNA2 (4963 nt; KM229701 = NC_035220) includes two potential ORFs (RNA2-ORF1 and RNA2-ORF2) (Sanfacon 2015; Seo et al. 2015; van der Vlugt et al. 2015; Thompson et al. 2017).

Pepper mild mottle virus Taxonomic position Genus: Tobamovirus

(PMMoV)

Family: Virgaviridae

PMMoV infection in plants of Leonurus sibiricus was reported from Korea (Kwon et al. 2016). The virus-infected honeyweed plants exhibit mild mottle-mosaic, and chlorosis symptoms. There is no known vector for this virus. The virus is transmissible by mechanical sap-inoculation, by contact between plants and also through seed. For more details of PMMoV, refer to Capsicum annuum.

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Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Leonurus sibiricus was reported from Korea (Kwon et al. 2016). The virusinfected honeyweed plants exhibit mild mottle, yellowing, and stunting symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow spot virus Taxonomic position Genus: Begomovirus

(ToYSV)

Family: Geminiviridae

ToYSV infection in plants of Leonurus sibiricus was reported from Paraguay and Brazil (Barbosa et al. 2013; Fernandes-Acioli et al. 2014; Ferro et al. 2017). The virus-infected honeyweed plants show yellow leaf mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The genome is a bipartite consisting of two circular, singlestranded DNA components. DNA-A consists of 2661 nt (KX348169) and DNA-B of 2629 nt (KX348214). A new alphasatellite is associated (Ferro et al. 2017). For more details of ToYSV, refer to Solanum lycopersicum.

References Barbosa JC, Eckstein B, Bergamin Filho A, Rezende JAM, Dallagnol LJ (2013) First report of Tomato yellow spot virus infecting Leonurus sibiricus in Brazil. Plant Dis 97:289 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer Index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Faria JC, Maxwell DP (1999) Variability in geminivirus isolates associated with Phaseolus spp. in Brazil. Phytopathology 89:262–268 Fernandes-Acioli NAN, Boiteux LS, Fonseca MEN, Segnana LRG, Kitajima EW (2014) Report of Tomato yellow spot virus infecting Leonurus sibiricus in Paraguay and within tomato fields in Brazil. Plant Dis 98:1445 Ferro CG, Silva JP, Xavier CAD, Godinho MT, Lima ATM, Mar TB, Lau D, Zerbini FM (2017) The ever increasing diversity of begomoviruses infecting non-cultivated hosts: new species from Sida spp. and Leonurus sibiricus, plus two New World alphasatellites. Ann Appl Biol 170:204–218 Kwon S-J, Choi G-S, Yoon J-Y, Seo J-K, Choi H-S (2016) Identification of Leonurus sibiricus as a weed reservoir for three pepper-infecting viruses. Plant Pathol J 32(1):65–69 Sanfacon H (2015) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. In: eLS. Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000764.pub3. http://www.els.net Seo J-K, Shin OJ, Kwak H-R, Kim M-K, Choi H-S, Lee S-H, Kim J-S (2014) First report of Broad bean wilt virus 2 in Leonurus sibiricus in Korea. Plant Dis 98:1748 Seo JK, Kang M, Kwak HR, Kim MK, Kim CS, Lee SH, Kim JS, Choi HS (2015) Complete genome sequence of motherwort yellow mottle virus, a novel putative member of the genus Torradovirus. Arch Virol 160:587–590 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J Gen Virol 98:529–531

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Lepidium spp.

van der Vlugt RAA, Verbeek M, Dullemans AM, Wintermantel WM, Cuellar WJ, Fox A, Thompson JR (2015) Torradoviruses. Annu Rev Phytopathol 53:485–512 Verbeek M, van Bekkum PJ, Dullemans AM, van der Vlugt RA (2014) Torradoviruses are transmitted in a semi-persistent and stylet-borne manner, by three whitefly vectors. Virus Res 186:55–60 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Lepidium spp. Family: Brassicaceae

Beet curly top virus Taxonomic position Genus: Curtovirus

Leafy vegetable

(BCTV)

Family: Geminiviridae

BCTV infection in plants of Lepidium spp. was reported from California, USA (Creamer et al. 1996). The virus is transmitted by a leafhopper vector, Circulifer tenellus, in a persistent (circulative, non-propagative) manner. The virus is not transmissible by mechanical inoculation. For more details of BCTV, refer to Beta vulgaris.

Bidens mottle virus Taxonomic position Genus: Potyvirus

(BiMoV)

Family: Potyviridae

BiMoV infection in plants of Lepidium virginicum was reported from Florida, USA (Christie et al. 1968). The virus-infected lepidium plants exhibit distortion of leaves, mild mottling, and stunting symptoms. The virus is transmitted by aphid vectors, Myzus persicae, Aphis spiraecola, Aphis craccivora, and Acyrthosiphon pisum, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BiMoV, refer to Bidens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Lepidium virginicum was reported from Georgia (USA) (Mullis et al. 2009). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. For more details of TSWV, refer to Solanum lycopersicum.

Lepistemon spp.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

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(TuMV)

Family: Potyviridae

TuMV infection in plants of Lepidium spp. was reported from New Zealand and China (Fletcher et al. 2009; Yin et al. 2015). The virus-infected lepidium plants showed interveinal yellow mottling, mosaic, distortion, and twisting of the leaves. The virus is transmitted by aphid species such as Myzus persicae and Brevicoryne brassicae in a non-persistent manner, and also by mechanical sap-inoculation to a large number of host plants. For more details of TuMV, refer to Brassica rapa.

References Christie SR, Edwardson JR, Zettler FW (1968) Characterization and electron microscopy of virus isolated from Bidens and Lepidium. Plant Dis Reptr 52:763–767 Creamer R, Lugue-Williams M, Howo M (1996) Epidemiology and incidence of Beet curly top geminivirus in naturally infected weed hosts. Plant Dis 80:533–535 Fletcher JD, Bulman S, Fletcher PJ, Houliston GJ (2009) First record of Turnip mosaic virus in Cook’s scurvy grass (Lepidium oleraceum agg.) – an endangered native plant in New Zealand. Aust Plant Dis 4:9–11 Mullis S, Bertrand PF, Brown SL, Csinos A, Diaz-Perez JC, Gitaitis R, Hickman LL, Johnson A, LaHue SS, Martinez N, McPherson RM, Nischwitz C, Reay-Jones F, Riley D, Sherwood J, Stevenson K, Wells L (2009) Tospoviruses in Solanaceae and other crops in the coastal plain of Georgia. University of Georgia, College of Agricultural and Environmental Sciences, Bulletin 1354, pp 51 Yin YY, Lu X, Li TT, He ZJ, Guo M, Ding M (2015) Detection of Turnip mosaic virus infecting Maca in Yunnan, Southwest of China. J Plant Pathol 97:S74

Lepistemon spp. Family: Convolvulaceae

Sweet potato feathery mottle virus Taxonomic position Genus: Potyvirus

(SPFMV)

Family: Potyviridae

SPFMV infection in plants of Lepistemon spp. was reported from Uganda (Tugume et al. 2010). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SPFMV, refer to Ipomoea batatas.

References Tugume AK, Mukasa SB, Valkonen JPT (2010) The significance of wild plants in the evolutionary diversification of Sweet potato feathery mottle virus in East Africa. In: 11th international plant virus epidemiology symposium held at Cornell University, New York, Abstract

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Lespedeza procumbens (Trailing lespedeza)

Lespedeza procumbens (Trailing lespedeza) Family: Fabaceae

Weed host

Trailing lespedeza virus 1 Taxonomic position Genus: Unassigned

(TLV-1)

Family: Tombusviridae

Geographical distribution TLV-1 infection in plants of Lespedeza procumbens was reported from the USA (Scheets et al. 2011). Symptoms and host(s) The virus-infected trailing lespedeza plants do not exhibit any symptoms. Virion properties and genome The virions are isometric and icosahedral (T = 3), non-enveloped, and 28–34 nm in diameter. The genome consists of a single molecule of positive-sense, linear, single-stranded RNA of 3929 nt (HM640935, NC_015227) (Scheets et al. 2011).

References Scheets K, Blinkova O, Melcher U, Palmer MW, Wiley GB, Ding T, Roe BA (2011) Detection of members of the Tombusviridae in the Tallgrass Prairie Preserve, Osage County, Oklahoma, USA. Virus Res 160(1–2): 256–263

Leucaena leucocephala (White lead tree) Family: Fabaceae

Trees/Shrubs

African cassava mosaic virus Taxonomic position Genus: Begomovirus

(ACMV)

Family: Geminiviridae

ACMV infection in trees of Leucaena leucocephala was reported from Nigeria (Alabi et al. 2008). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of ACMV, refer to Manihot esculenta.

Leucanthemum vulgare (Oxeye daisy)

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East African cassava mosaic Cameroon virus Taxonomic position Genus: Begomovirus

(EACMCV)

Family: Geminiviridae

ACMV infection in trees of Leucaena leucocephala was reported from Nigeria (Alabi et al. 2008). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of EACMCV, refer to Manihot esculenta.

References Alabi OJ, Ogbe FO, Bandyopadhyay R, Lava Kumar P, Dixon AGO, Hughes J’A, Naidu RA (2008) Alternate hosts of African cassava mosaic virus and East African cassava mosaic Cameroon virus in Nigeria. Arch Virol 153:1743–1747

Leucanthemum vulgare (Oxeye daisy) Family: Asteraceae

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

L

Ornamental

(INSV)

Family: Tospoviridae

INSV infection was identified in plants of Leucanthemum spp. in Iran (Shahraeen et al. 2002). The virus-infected oxeye daisy plants exhibit symptoms of mottling/mosaic, foliar distortion and chlorosis, and ring-spots. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Potato yellow dwarf nucleorhabdovirus Taxonomic position Genus: Nucleorhabdovirus

(PYDV)

Family: Rhabdoviridae

PYDV infection in plants of Leucanthemum vulgare was reported from California, USA (Younkin 1942). The virus-infected oxeye daisy plants initially showed vein-clearing, followed by pale green veins on plants with a rosette of stunted foliage and shortened petioles. The virus is transmitted by leafhopper vectors in a persistent, propagative manner, and also by mechanical sap-inoculation. For more details of PYDV, refer to Solanum tuberosum.

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Leucocoryne purpurea (Glory-of-the-sun)

References Shahraeen N, Ghotbi T, Mehraban AH (2002) Occurrence of Impatiens necrotic spot virus in ornamentals in Mahallat and Tehran Provinces in Iran. Plant Dis 86:694 Younkin SG (1942) Weed suspects of the Potato yellow dwarf virus. Am Potato J 19:6–11

Leucocoryne purpurea (Glory-of-the-sun) Family: Amaryllidaceae

Ornamental

Turnip mosaic virus

(TuMV)

Taxonomic position Genus: Potyvirus

Family: Potyviridae

TuMV infection in plants of Leucocoryne purpurea was reported from Chile (Camps et al. 2016). The virus-infected leucocoryne plants exhibit symptoms such as violet spots on the leaves, curling of the stems, and off-color mosaic patterns on the petals. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of host plants. For more details of TuMV, refer to Brassica rapa.

References Camps R, Mansur L, De La Cuadra C, Besoain X (2016) First report of Turnip mosaic virus (TuMV) infecting Leucocoryne purpurea (Amaryllidaceae) in Chile. Plant Dis 100:2341

Liatris spicata (Blazing star and gayfeather) Family: Asteraceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

Family: Secoviridae

ArMV infection in plants of Liatris spicata was reported from Lithuania (Samuitiene et al. 2008). The virus-infected blazing star plants showed light green spots and mosaic symptoms. The virus is transmitted by the nematode vectors, Xiphinema diversicaudatum and X. coxi, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Liatris spicata (Blazing star and gayfeather)

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

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(BYMV)

Family: Potyviridae

BYMV infection in plants of Liatris spp. was reported from South Africa (Schulze et al. 2017). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Liatris spicata was reported from the Netherlands (Derks et al. 1986). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Gayfeather mild mottle virus Taxonomic position Genus: Cucumovirus

(GMMV)

L Family: Bromoviridae

Geographical distribution GMMV infection in plants of Liatris spicata was reported from Poland (Adams et al. 2009). Symptoms and host(s) The virus-infected gayfeather plants showed mild mottle symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner, and the virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped and spherical, of about 25–30 nm in diameter with T = 3 icosahedral symmetry, and composed of 180 coat proteins: 12 pentamers and 20 hexamers. The genome is a segmented, tripartite, linear, positive-sense ssRNA composed of RNA1 of 3350 nt (FM881899 = NC_012134), RNA2 of 2935 nt (FM881900 = NC_012135), and RNA3 of 2214 nt (FM881901 = NC_012136). Each genomic segment has a 30 tRNA-like structure and a 50 cap (Garcia-Arenal Rodriguez and Fraile 2011; Scott 2011).

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

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Ligustrum spp. (Privet)

ToRSV was reported from plants of Liatris spicata in Lithuania (Navalinskien_e and Samuitien_e 2006). The virus-infected blazing star plants exhibit symptoms of malformation of leaves and flowers, chlorotic or necrotic spots, streaks, or ring-spots and veinal necrosis. The virus is transmitted by nematode vectors (Xiphinema spp.) in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and also by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

References Adams IP, Glover RH, Monger WA, Mumford R, Jackeviciene E, Navalinskiene M, Samuitiene M, Boonham N (2009) Next-generation sequencing and metagenomic analysis: a universal diagnostic tool in plant virology. Mol Plant Pathol 10:537–545 Derks AFLM, Hollinger TC, Segers LC, Asjes CJ (1986) In: Department of Plant Pathology. Jaarverslag 1985, Laboratorium voor Bloembollenonderzoek, Lisse. pp 55–78 Garcia-Arenal Rodriguez F, Fraile A (2011) Cucumovirus. Bromoviridae. In: The Springer Index of Viruses. Springer, New York, pp 179–185. https://doi.org/10.1007/978-0-387-95919-1_26 Navalinskien_e M, Samuitien_e M (2006) Dekoratyvinių augalų virusin_es ligos ir jų suk_el_ejai Lietuvoje. Lutut_e, Kaunas 254 p Samuitiene M, Navalinskiene M, Jackeviciene E (2008) Arabis mosaic virus on ornamental plants. Biologia 54:264–268 Schulze A, Roberts R, Pietersen G (2017) First report of the detection of Bean yellow mosaic virus (BYMV) on Tropaeolum majus; Hippeastrum spp., and Liatris spp. in South Africa. Plant Dis 101:846 Scott SW (2011) Bromoviridae and allies. In: Encyclopedia of Life Sciences (ELS). Wiley, Chichester. https://doi.org/ 10.1002/9780470015902

Ligustrum spp. (Privet) Family: Oleaceae

Arabis mosaic virus Taxonomic position Genus: Nepovirus

Ornamental

(ArMV)

Family: Secoviridae

ArMV infection in plants of Ligustrum spp. was reported from the UK, Germany, and Bulgaria (Cadman 1960; Schmelzer 1963; Jankulova and Schmelzer 1974; Dupuis et al. 2008), and a similar disease has been reported in the USA (Plakidas 1959; Burnett and Youtsey 1962) in multiple species of ligustrum. Symptoms are described as bright yellow rings and spots with chlorotic rings and sometimes depressed or necrotic lesion (but note that similar symptoms occur in plants infected with Ligustrum necrotic ringspot virus, and earlier reports from the USA do not identify the causal agent beyond “virus”). This virus is transmitted by nematode vectors (Xiphinema spp.) in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Cherry leaf roll virus Taxonomic position Genus: Nepovirus

(CLRV)

Family: Secoviridae

Ligustrum spp. (Privet)

1371

CLRV-infected plants of Ligustrum spp. were reported from Germany (Schmelzer 1972). The virusinfected privet plants exhibit ring-spot symptoms. The virus is transmitted by nematode vectors Xiphinema coxi, X. diversicaudatum, and X. vuittenezi in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants. For more details of CLRV, refer to Prunus avium.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Ligustrum spp. was reported from the Europe, Poland, and North America (Smith 1957; Kochman et al. 1964). The virus-infected privet plants exhibit symptoms of chlorotic chevrons, rings, or blotches. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Ligustrum necrotic ringspot virus Taxonomic position Genus: Carlavirus

(LNRSV)

L Family: Betaflexiviridae

Geographical distribution LNRSV infection in plants of Ligustrum japonicum was reported from the USA (Burnett and Youtsey 1962; Scott and Zimmerman 2008).

Symptoms and host(s) The virus-infected privet plants show necrotic ring-spots and line patterns. LNRSV has also been reported from Mazus reptans, Impatiens spp., and Phlox spp.

Transmission The virus was shown to be graft-transmissible. The virus expected to be transmitted by aphid vectors in a non-persistent manner, and is also mechanically sap transmissible. The virus is mainly disseminated through the use of virus-infected vegetative propagative materials.

Virion properties and genome The virions are flexuous filaments about 650 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8412 nt (EU074853 = NC_010305) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Adams et al. 2004; Scott and Zimmerman 2008).

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Ligustrum spp. (Privet)

Ligustrum ringspot virus

(LigRSV)

Taxonomic position LigRSV is a tentative member of the genus Cilevirus of family Kitaviridae. Geographical distribution LigRSV infection in plants of Ligustrum lucidum and L. sinensis has been reported from Argentina and Brazil (Da Costa Lima et al. 1991; Kitajima et al. 2003, 2010). Symptoms and host(s) The virus-infected privet plants exhibit symptoms of chlorotic spots or rings. Transmission The virus is transmitted by mite vectors, Brevipalpus obovatus (syn. Tenuipalpus pseudocuneatus) and B. phoenicis. Virion properties and genome The virus induces electron-dense viroplasms in the cytoplasm of mesophyll cells within the lesions, and short bacilliform particles may also be observed; no specific description of particle properties is provided. The genome is bipartite, of two single-stranded, polyadenylated RNAs. Partial genomic sequences are available (HM164551, 714 nt; HM164552, 1591 nt).

Ligustrum virus A Taxonomic position Genus: Carlavirus

(LiVA)

Family: Betaflexiviridae

Geographical distribution LiVA infection in plants of Ligustrum obtusifolium was reported from South Korea (Igori et al. 2016). Transmission The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. Virion properties and genome Virions are flexuous filaments about 580 nm in length and 13 nm in diameter with helical symmetry. The genome contains a single molecule of linear ssRNA of 8525 nt (KX000914 = NC_031089), excluding the poly(A) tail, and consists of six open reading frames (Igori et al. 2016).

Petunia asteroid mosaic virus Taxonomic position Genus: Tombusvirus

(PetAMV)

Family: Tombusviridae

Ligustrum spp. (Privet)

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PetAMV was reported from the roots, but not leaves, of Ligustrum vulgare in Italy (Lovisolo et al. 1965). The virus-infected privet plants showed some chlorotic leaf spotting. The virus is transmissible by mechanical sap-inoculation, and also by grafting. No vector is reported for this virus. For more details of PetAMV, refer to Petunia spp.

Plum pox virus Taxonomic position Genus: Potyvirus

(PPV)

Family: Potyviridae

PPV infection in plants of Ligustrum vulgare was reported from the Czech Republic (Polak 2001). The virus-infected privet plants exhibit symptoms of diffuse spots and rings. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PPV, refer to Prunus domestica.

Privet idaeovirus Synonyms Privet leaf blotch-associated virus (PrLBaV) Taxonomic position Genus: Idaeovirus

L Family: Unassigned

Geographical distribution The virus infection in plants of Ligustrum japonicum was reported from Italy (Navarro et al. 2017). Symptoms and host(s) The virus-infected privet plants exhibit leaf blotch symptoms. Transmission The virus is transmitted by mechanical sap inoculation. The virus is graft transmissible. The use of virus-infected propagative material will further spread the disease. Virion properties and genome The virions are isometric (icosahedral) but slightly flattened, not enveloped, of three types but similar in size (33 nm). The genome composed of two positive single-stranded RNAs, one of which, RNA1 consisting of 5377 nt (LT221868 = NC_031341), is monocistronic and codes for the viral replicase, whereas the other RNA2 is 2348 nt (LT221869 = NC_031342), and contains two open reading frames (ORFs), ORF2a and ORF2b, coding for the putative movement (p38) and coat (p30) proteins (Navarro et al. 2017).

Privet ringspot virus (PrRSV) Taxonomic position Genus: Ilarvirus

Family:Bromoviridae

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Ligustrum spp. (Privet)

Geographical distribution PrRSV infection in plants of Ligustrum japonicum was reported from the USA (Burnett and Youtsey 1962; Aboughanem-Sabanadzovic et al. 2016). Symptoms and host(s) The virus-infected privet plants exhibit ringspots, irregular line patterns and foliar deformation symptoms. Transmission The virus is transmitted by different thrips species. Infected pollen is blown to healthy plants where thrips feeding on the pollen mechanically transmit the virus. The virus is carried both inside the pollen and on the surface. Virus is transmitted by mechanical sap-inoculation, and by grafting. The virus is not transmitted by contact between plants. Virion properties and genome The virus has non-enveloped isometric and quasi-isometric (bacilliform) particles of 23, 25, and 27 nm. The genome is segmented, tripartite linear positive-sense ssRNA composed of RNA1 (3415 nt; KT290039 = NC_027928); RNA 2 (2863 nt; KT290040 = NC_027929); and RNA 3 (2140 nt; KT290041 = NC_027930) (Scott 2011; Aboughanem-Sabanadzovic et al. 2016).

Raspberry ringspot virus Taxonomic position Genus: Nepovirus

(RpRSV)

Family: Secoviridae

RpRSV infection in plants of Ligustrum spp. was reported from Denmark (Thomsen 1970). The virusinfected privet plants do not exhibit symptoms. The virus is transmitted by nematode vectors (Longidorus spp.) in a non-persistent manner, and also by mechanical sap-inoculation. For more details of RpRSV, refer to Rubus spp.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV infection in plants of Ligustrum spp. was reported from Europe (Thomsen 1970; Albouy and Devergne 1999). The virus-infected privet plants do not exhibit symptoms. There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco necrosis virus

(TNV)

Taxonomic position TNV is a tentative member of the genus Necrovirus and family Tombusviridae

Ligustrum spp. (Privet)

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TNV infection in plants of Ligustrum vulgare was reported from Europe (Paulechova and Baumgartnerova 1989; Albouy and Devergne 1999). The virus-infected privet plants exhibit mosaic symptoms. The virus is transmitted by the zoospores of the fungus Olpidium brassicae, and the virus is also mechanically sap-transmissible. It is not clear whether the TNV in ligustrum is TNV-A (Alphanecrovirus) or TNV-D (Betanecrovirus). For more details of TNV, refer to Nicotiana tabacum.

Tomato black ring virus Taxonomic position Genus: Nepovirus

(TBRV)

Family: Secoviridae

TBRV infection in plants of Ligustrum vulgare was reported from Poland (Blaszczak and Pospieszny 1987). The virus is transmitted by the nematode vector Longidorus attenuatus in a non-persistent manner and also by mechanical sap-inoculation. For more details of TBRV, refer to Solanum lycopersicum.

Tomato bushy stunt virus Taxonomic position Genus: Tombusvirus

(TBSV)

Family: Tombusviridae

TBSV infection in plants of Ligustrum vulgare was reported by Novak and Lanzova (1980). The virus is transmissible by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants. No known vector is reported for this virus. For more details of TBSV, refer to Solanum lycopersicum.

References Aboughanem-Sabanadzovic N, Tzanetakis IE, Lawrence A, Stephenson RC, Sabanadzovic S (2016) A novel Ilarvirus is associated with Privet necrotic ringspot disease in the Southern United States. Phytopathology 106:87–93 Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Albouy J, Devergne JC (1999) Diseases produced by viruses on ornamental plants. Ediciones Mundi-Prensa, Madrid 480 p Blaszczak W, Pospieszny H (1987) Ligustrum vulgare L. – a natural host of the Tomato black ring virus. Pr Nauk Inst Ochr Rosl 29:127–136 Burnett HC, Youtsey CO (1962) Necrotic ringspot, a virus disease of Ligustrum spp. Fla State Hortic Soc 1962:472–476 Cadman CH (1960) Viruses with nematodes as vectors. Rep Scott Hortic Res Inst 1960–1:54 Da Costa Lima MLRZ, Lim Neto VC, de Souza VBV (1991) The causal agent of ligustrum ringspot disease. Phytopathology 81:1216 Dupuis L, Cobanov P, Bassler A, Krczal G, Wetzel T (2008) Complete genome sequence of a virulent isolate of Arabis mosaic virus from privet (Ligustrum vulgare). Arch Virol 153:1611–1613 Igori D, Lim S, Zhao F, Baek D, Park JM, Cho HS, Kim HS, Kwon SY, Moon JS (2016) The complete sequence and genome organization of Ligustrum virus A, a novel Carlavirus. Arch Virol 161:3593–3596 Jankulova M, Schmelzer K (1974) Esrtmaliger nachweis des Arabis – mosaik-virus in der Volksrepublik Bulgarien. Arch Phytopathol Pflanzenschutz 10:283–284 Kitajima EW, Chagas CM, Rodrigues JCV (2003) Brevipalpus-transmitted plant virus and virus-like diseases: cytopathology and reports of some recent cases. Exp Appl Acarol 30:135–160

L

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Lilium spp. (Lilium longiflorum, L. lancifolium, L. tsingtauense)

Kitajima EW, Rodrigues JCV, Freitas-Astúa J (2010) An annotated list of ornamentals naturally found infected by Brevipalpus mite-transmitted viruses. Sci Agric 67:348–371 Kochman J, Kowalska H, Syzmanska B (1964) Untersuchungen uber virosen des flieders (Syringa vulgaris L.) und des ligusters (Ligustrum vulgare L.). Phytopathol Z 51:333–350 Lovisolo O, Bode O, Volk J (1965) Preliminary studies on the soil transmission of Petunia asteroid mosaic virus (= ‘Petunia’ strain of tomato bushy stunt virus). Phytopathol Z 53:323–342 Navarro B, Loconsole G, Giampetruzzi A, Aboughanem-Sabanadzovic N, Ragozzino A, Ragozzino E, Di Serio F (2017) Identification and characterization of Privet leaf blotch-associated virus, a novel Idaeovirus. Mol Plant Pathol 18(7):925–936 Novak JB, Lanzova J (1980) Demonstration of tomato bushy stunt virus in some forest tree species and plants. Lesnictví 26:1009–1016 Paulechova K, Baumgartnerova H (1989) Some properties of the tobacco necrosis virus isolated from common privet ligustrum vulgare. Sbornik Uvtiz (Ustav Vedeckotechnickych Informaci Pro Zemedelstvi) Ochrana Rostlin 25(3):165–170 Plakidas AG (1959) Chlorotic spot, a graft-transmissible disease of Ligustrum. Plant Dis Reptr 43:688–689 Polak J (2001) European spindle tree and common privet, new natural hosts of Plum pox virus. Acta Hortic 550:125–128 Schmelzer K (1963) Investigations on viruses of ornamental and wild woody plants, IInd part. Viruses of forsythia, lonicera, ligustrum and laburnum. Phytopathol Z 46:105–138 Schmelzer K (1972) Das Kirschenblattrollvirus aus der Birke (Betula pendula Roth.). Zentbl Bakteriol Z Abt II 127:10–12 Scott SW (2011) Ilarvirus. Bromoviridae. The Springer index of viruses. Springer, New York, pp 187–194. https://doi. org/10.1007/978-0-387-95919-1_27 Scott SW, Zimmerman MT (2008) The complete sequence of ligustrum necrotic ringspot virus, a novel carlavirus. Arch Virol 153:393–396 Smith KM (1957) A text book of plant virus diseases, 2nd edn. Churchill, London 572 pp Thomsen A (1970) Virus infection of Ligustrum spp. Tidssk PlAvl 74:234–242

Lilium spp. (Lilium longiflorum, L. lancifolium, L. tsingtauense) Family: Liliaceae

Ornamental

Apple stem grooving virus

(ASGV)

Synonyms Citrus tatter leaf virus Taxonomic position Genus: Capillovirus

Family: Betaflexiviridae

ASGV infection in plants of Lilium spp. was reported from Japan (Inouye et al. 1979). The virus was shown to cause lily yellowing disease; other isolates infect citrus, apple, and pear. No vector has been identified for this virus. The virus is mechanically sap-transmissible. The lily isolate was found to be seed-transmitted in lily, but not to be transmitted by aphids. For more details of ASGV, refer to Malus domestica.

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

Lilium spp. (Lilium longiflorum, L. lancifolium, L. tsingtauense)

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ArMV infection in plants of Lilium spp. was reported from the Netherlands, China, and Lithuania (Asjes and Segers 1983; Samuitiene et al. 2008). The virus-infected lily plants exhibit symptoms of necrosis, stunting, and death of the plants. This virus is transmitted by nematode vectors (Xiphinema spp.) in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Broad bean wilt virus Taxonomic position Genus: Fabavirus

(BBWV)

Family: Secoviridae

BBWV (but without specifying either of the currently recognized species BBWV-1 or BBWV-2) infection in plants of Lilium longiflorum was reported from Korea (Lee et al. 1996). The virus-infected lily plants exhibit symptoms of dwarfing, malformation, and color breaking. The virus is transmitted by an aphid vector, Myzus persicae, in a non-persistent manner, and also by mechanical sap-inoculation. It is not clear whether the lily isolate(s) were BBWV-1 or BBWV-2, but BBWV-2 has been documented in other crops in Korea. For more details of BBWV, refer to Vicia faba.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

L Family: Bromoviridae

CMV infection in plants of Lilium spp. was reported from Israel, India, Lithuania, Japan, Korea, China, and Poland (Benetti and Tomassoli 1988; Chang and Pang 1991; Chang et al. 1996; Cohen et al. 1996; Lee et al. 1996; Ram et al. 1999; Jung et al. 2000; Navalinskiene and Samuitiene 2001; Bellardi et al. 2002; Masuta et al. 2002; Ryu et al. 2002; Niimi et al. 2003; Sharma et al. 2005; Samuitiene and Navalinskiene 2008; Berniak et al. 2010; Wang et al. 2010). The virus-infected lilium plants show chlorotic or yellow spots or stripes or vein-clearing. Later gray or brown necrotic spots/stripes may develop. Leaves and petals may curl or may be malformed. This virus has a wide range of host plants, including many common weeds and cultivated plants. The leaves may become pale and the plant stunted. This virus is spread by aphids in a non-persistent manner (Hagita 1989). The use of scales from the virus-infected bulbs for vegetative reproduction is one of the major causes of virus spread. The virus is also mechanically sap-transmissible to a large number of host plants. For more details of CMV, refer to Cucumis sativus.

Iris yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(IYSV)

Family: Tospoviridae

IYSV was reported in plants of Lilium spp. in Israel (Gera et al. 2008). The virus-infected lilium plants exhibited necrotic spots and rings on the leaves and stunting. The virus is transmitted by a onion thrips vector, Thrips tabaci in a persistent-propagative manner and also by mechanical sap-inoculation. Use of scales from virus infected bulbs for vegetative reproduction is one of the major causes of virus spread. For more details of IYSV, refer to Iris spp. or Allium cepa.

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Lily latent virus

Lilium spp. (Lilium longiflorum, L. lancifolium, L. tsingtauense)

(LiLV)

Taxonomic position LiLV is a tentative member of the genus Carlavirus and family Betaflexiviridae. Geographical distribution LiLV infection in plants of Lilium lancifolium was reported from Korea (Ryu et al. 2000). Symptoms and host(s) The virus was detected in the Korean native lily, Lilium lancifolium, showing very mild mosaic or no apparent symptoms. Transmission The virus is presumed to be transmitted by aphids. Virion properties and genome The virions are filaments, c. 680  12 nm, encapsidating a single-stranded RNA of c.8.0 Kb. A partial genome sequence of 3000 nt is available (AJ131812) and has c.72–79% identity to multiple isolates of Lily symptomless virus, indicating that LiLV is a distinct species.

Lily mottle virus

(LMoV)

Synonyms Tulip band breaking virus (TBBV) Taxonomic position Genus: Potyvirus

Family: Potyviridae

Geographical distribution LMoV infection in plants of Lilium spp. was reported from India, Japan, Germany, Israel, Italy, the Netherlands, Lithuania, China, the UK, and the USA (Alper et al. 1982; Maeda et al. 1984; van Schadewijk 1986; Derks et al. 1994; Chang et al. 1996; Cohen et al. 1996; Miyagawa 1997; Asjes 2000; Navalinskiene and Samuitiene 2001; Niimi et al. 2003; Sharma et al. 2005; Wang et al. 2010; Rizzo et al. 2012; Fisher et al. 2013; Aravintharaj et al. 2017). Many reports of TBV from lily actually refer to Lily mottle virus (Yamaji et al. 2001). Symptoms and host(s) The virus-infected lilium leaves show a light to yellowish green mottle or mosaic and can be twisted or narrowed. Flowers are sometimes malformed or may show a breaking pattern. Bulb scales of some cultivars show brown necrotic spots or rings. The infected plants are often stunted and die prematurely (Mokra 1976) also causing vein-clearing, leaf mottle or mosaic, and reddish brown necrotic spots on Asiatic hybrid lilies (Rizzo et al. 2012). Transmission The virus is transmitted by aphid vectors Aphis gossypii, Myzus persicae, and Macrosiphum solanifolii in a non-persistent manner. The virus is sap-transmissible, and susceptible host species are found in the

Lilium spp. (Lilium longiflorum, L. lancifolium, L. tsingtauense)

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family Liliaceae. The virus is not seed-borne. The use of scales from the virus-infected bulbs for vegetative reproduction is one of the major causes of virus spread. Virion properties and genome The virions are non-enveloped, flexuous filaments, 750–770 nm long, and 11–13 nm wide. The genome consists of a single molecule of a linear, positive-sense ssRNA of 9644 nt (AJ564636 = NC_005288) (Zheng et al. 2003; Revers and Garcia 2015; Wylie et al. 2017).

Lily symptomless virus

(LSV)

Synonyms Lily rosette virus Taxonomic position Genus: Carlavirus

Family: Betaflexiviridae

Geographical distribution LSVoccurs wherever Lilium spp. are grown including the USA, Israel, India, Lithuania, China, and the Netherlands (Brierley and Smith 1945; Allen et al. 1980; van Schadewijk 1986; Hsu et al. 1995; Blake and Wilson 1996; Chang et al. 1996; Cohen et al. 1996; Asjes 2000; Bellardi et al. 2002; Niimi et al. 2003; Sharma et al. 2005; Wang et al. 2010; Fisher 2013; Fisher et al. 2013). Symptoms and host(s) The virus-infected lilies exhibit symptoms such as yellowing, chlorotic striping between the veins, vein-clearing or light green stripes between the veins, and reduced growth of the plants. Severely infected lily plants produced smaller deformed flowers and significantly lower bulb yields. In some lily cultivars, yellow, purple, or brown spots develop on the upper surface of the leaves which leads to premature dieback of the plants at the end of the growing season (Allen 1972; Fisher 2013). Transmission The virus is transmitted by aphid species: including Aphis gossypii, A. fabae, Aulacorthum solani, and Macrosiphum euphorbiae in a non-persistent manner (Mowat and Stefanac 1974). This virus is mechanically sap-transmissible to the members of the family Liliaceae, Chenopodiaceae, and Solanaceae, and the virus is not seed-borne in lilies. The use of scales from the virus-infected bulbs for vegetative reproduction is one of the major causes of virus spread. Virion properties and genome The virions are flexuous filaments about 640 nm in length and 17–18 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8394 nt (AJ516059 = NC_005138) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Derks and Vink-Van den Abeele 1980; Hagita 1989; Choi and Ryu 2003; Zheng et al. 2003; Adams et al. 2004; Singh et al. 2005).

L

1380

Lily virus A

Lilium spp. (Lilium longiflorum, L. lancifolium, L. tsingtauense)

(LVA)

Taxonomic position LVA is a tentative member of the genus Potyvirus and family Potyviridae Geographical distribution LVA infection in plants of Lilium longiflorum was reported from Australia (Wylie et al. 2012). Transmission The virus is presumed to be transmitted by aphid vectors. Virion properties and genome The virions are flexuous filamentous. The genome consists of a single molecule of positive-sense singlestranded RNA. A partial polyprotein gene sequence of 7900 nt is available (JN127335) (Wylie et al. 2017).

Lily virus X

(LVX)

Taxonomic position Genus: Potexvirus

Family: Alphaflexiviridae

Geographical distribution LVX infection in plants of Lilium spp. occurs in the UK, the Netherlands, and Israel (Derks and Hendriks 1990; Cohen et al. 1996; Yang et al. 1996). Symptoms and host(s) The virus was originally detected in symptomless plants of Lilium formosanum. The virus-infected lily plants may show faint chlorotic spots on the leaves and occasionally brown necrotic lesions. LVX has also been reported in Tricyrtis formosana (toad lily) in the USA. Transmission The virus is transmissible by mechanical sap-inoculation, and susceptible host species are found in the families Amaranthaceae, Chenopodiaceae, Liliaceae, Solanaceae, and Aizoaceae. The virus is not transmissible by contact between plants. No vector is identified (Stone 1980). The use of scales from the virus-infected bulbs for vegetative reproduction is one of the major causes of virus spread. Virion properties and Genome The virions are flexuous, filaments, c.470 nm long, and 13 nm wide. The genome is a single linear molecule of positive-sense, single-stranded RNA of 5823 nt (AJ633822 = NC_007192) and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type 18–27 kDa in size (Adams et al. 2004; Chen et al. 2005).

Narcissus mosaic virus Taxonomic position Genus: Potexvirus

(NMV)

Family: Alphaflexiviridae

Lilium spp. (Lilium longiflorum, L. lancifolium, L. tsingtauense)

1381

NMV was reported affecting plants of Lilium speciosum in Italy (Bellardi et al. 1988). The virusinfected lilium plants showed severe symptoms including yellow stripes on leaves, leaf deformation and stunting, and in some cases premature plant death. The virus is mechanically sap-transmissible. For more details of NMV, refer to Narcissus spp.

Plantago asiatica mosaic virus Taxonomic position Genus: Potexvirus

(PlAMV)

Family: Alphaflexiviridae

PlAMV infection in plants of Lilium spp. was reported from Japan, the Netherlands, the USA, Hungary, Taiwan, Italy, Korea, Costa Rica, China, and Chile (Komatsu et al. 2008; Anonymous 2010; Chen et al. 2013; Hammond et al. 2015; Pajtli et al. 2015; Parrella et al. 2015; Vidal et al. 2016; Montero-Astua et al. 2017; Xu et al. 2017). The virus-infected plants of Oriental and Asiatic lily hybrids may show chlorotic and necrotic streaking on leaves and necrotic streaking on stems, becoming most apparent after flowering. Transmission appears to occur through the planting medium, but no vector is known. The virus is mechanically sap-transmissible. For more details of PlAMV, refer to Plantago asiatica.

Rembrandt tulip breaking virus

(ReTBV)

Taxonomic position ReTBV is a tentative member of the genus Potyvirus and family Potyviridae ReTBV infection in plants of Asiatic hybrid lily was reported from Japan (Yamamoto and Senda 2012). The virus-infected lily plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors, and the virus is also mechanically sap-transmissible. For more details of ReTBV, refer to Tulipa spp.

Strawberry latent ringspot virus Taxonomic position Genus: Unassigned

(SLRSV)

Family: Secoviridae

SLRSV infection was reported in plants of Oriental lily hybrids in India, Israel, and Italy (Cohen et al. 1995, 1996; Bellardi et al. 2002; Sharma et al. 2005). The virus-infected lily plants exhibit strong foliar mosaic and asymmetrical opening of flowers. The virus is transmitted by nematode vector (Xiphinema spp.) and also by mechanical sap-inoculation. The virus is transmissible by grafting. For more details of SLRSV, refer to Fragaria spp.

Tobacco necrosis virus

(TNV)

Taxonomic position TNV is a tentative member of the genus Necrovirus and family Tombusviridae TNV infection in plants of Lilium spp. was reported by Derks et al. (2003). The virus-infected lily plants exhibit symptoms of necrotic streaks in the foliage. The virus is transmitted by the zoospores of the

L

1382

Lilium spp. (Lilium longiflorum, L. lancifolium, L. tsingtauense)

fungus Olpidium brassicae, and the virus is also mechanically sap-transmissible. It is not clear whether the lily isolate(s) of TNV are of the TNV-A (Alphanecrovirus) or TNV-D (Betanecrovirus). For more details of TNV, refer to Nicotiana tabacum.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in plants of Lilium spp. was reported from Italy and Lithuania (Derks 1975; Tomassoli and Bennetti 1988; Navalinskiene and Samuitiene 2001). The virus-infected lily plants showed chlorotic or necrotic mosaic, together with distortion and down curling of leaves. The virus is transmitted by nematode vectors, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Lilium longiflorum was reported from the USA (Travis and Brierley 1957). No symptoms were observed in naturally infected lily plants. The virus is transmitted by nematode vectors in a non-persistent manner. The virus is mechanically sap-transmissible and also through grafting. The use of vegetative propagating material from virus-infected plants is the primary mode of virus spread. For more details of TRSV, refer to Nicotiana tabacum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Lilium spp. was reported from Korea, Italy, and Lithuania (Kim and Choi 1990; Lee et al. 1996; Navalinskiene and Samuitiene 2001; Bellardi et al. 2002). The virus-infected lily plants exhibit distorted leaves with chlorotic spots and streak symptoms. The virus is transmitted by a nematode vector (Xiphinema spp.) in a non-persistent manner. The virus is transmissible by mechanical sap-inoculation, and also by grafting. For more details of ToRSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Lilium spp. was reported from Lithuania (Navalinskiene and Samuitiene 2001; Parrella et al. 2003). The virus-infected lily plants exhibit symptoms of stunting and leaves which

Lilium spp. (Lilium longiflorum, L. lancifolium, L. tsingtauense)

1383

are distorted, with chlorotic and necrotic spots. The virus is transmitted by thrips vectors in a persistentpropagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Turnip mosaic virus

(TuMV)

Synonyms Tulip top-breaking virus Taxonomic position Genus: Potyvirus

Family: Potyviridae

TuMV infection in pink-flowered lily plants was reported from Italy (Bellardi et al. 2002). The virus-infected lily plants exhibit flower break symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

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Limonium spp. (Statice)

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Ryu JH, Park HW, Park WM, Lee SY, Ryu KH (2000) Molecular analysis of the 30 terminal region of Lily latent carlavirus from Lilium lancifolium. Plant Pathol J 16:231–235 Ryu KH, Park HW, Choi JK (2002) Characterization and sequence analysis of a lily isolate of Cucumber mosaic virus from Lilium tsingtauense. Plant Pathol J 18:85–92 Samuitiene M, Navalinskiene M (2008) Occurrence of Cucumber mosaic cucumovirus on ornamental plants in Lithuania. Zemdirbyste-Agriculture 95:135–143 Samuitiene M, Navalinskiene M, Jackeviciene E (2008) Arabis mosaic virus on ornamental plants. Biologia 54:264–268 Sharma A, Mahinghara BK, Singh AK, Kulshrestha S, Raikhy G, Singh L, Verma N, Hallan V, Ram R, Zaidi AA (2005) Identification, detection and frequency of lily viruses in Northern India. Scientia Hortic 106:213–227 Singh AK, Hallan V, Ram R, Zaidi AA (2005) Variability in the coat protein of Lily symptomless virus isolates infecting various lily species. Plant Pathol 54:621–624 Stone OM (1980) Two new potexviruses from monocotyledons. Acta Hortic 110:59–63 Tomassoli L, Bennetti MP (1988) Tobacco rattle virus in triple infection in lily. Adv Hortic Sci 2:117–119 Travis RV, Brierley P (1957) Tobacco ring spot virus from iris and Easter lily. Plant Dis Reptr 41:254 van Schadewijk AR (1986) Detection of Tulip breaking virus and Lily symptomless virus by means of ELISA. Acta Hortic 177:121–128 Vidal AK, Camps R, Besoian XA (2016) First report of necrotic streaking of Asiatic lilies caused by Plantago asiatica mosaic virus in Chile. Plant Dis 100:1799 Wang R, Wang G, Zhao Q, Zhu Y, Zhan J, Xie Z, An L, Wang Y (2010) Molecular and cytopathologic evidences for a mixed infection of multiple viruses on Lanzhou lily (Lilium davidii var. unicolor) in Northwestern China. J Plant Dis Protect 117:145–149 Wylie SJ, Luo H, Li H, Jones MG (2012) Multiple polyadenylated RNA viruses detected in pooled cultivated and wild plant samples. Arch Virol 157(2):271–284 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Xu LF, Ming J, Yuan SX (2017) First report of Plantago asiatica mosaic virus in lily hybrids in China. Plant Dis 101:263 Yamaji Y, Lu X, Kagiwada S, Ohshima K, Nama S (2001) Molecular evidence that a liy-infecting strain of Tulip breaking virus from Japan is a strain of Lily Mottle virus. Eur J Plant Pathol 107:833–837 Yamamoto H, Senda M (2012) Rembrandt tulip breaking virus detected from lily. Jpn J Phytopathol 78:111–113 Yang TC, Zettller FW, Polston JE (1996) First report of Lily X potexvirus in the United States. Plant Dis 80:1430 Zheng HY, Chen J, Zhao MF, Lin L, Chen JP, Antoniw JF, Adams MJ (2003) Occurrence and sequences of Lily mottle virus and Lily symptomless virus in plants grown from imported bulbs in Zhejiang province. China Arch Virol 148(12):2419–2428

Limonium spp. (Statice) Family: Plumbaginaceae

Broad bean wilt virus Taxonomic position Genus: Fabavirus

Ornamental

(BBWV)

Family: Secoviridae

BBWV (but without specifying either of the currently recognized species BBWV-1 or BBWV-2) infection in plants of Limonium sinuatum was first reported from Southern Germany (Hein et al. 1977; Matsumoto et al. 1997). The virus-infected statice plants had small puckered, twisted, wilted, and deformed leaves, and some of the leaves exhibited a mosaic pattern; the infected plants were stunted. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a number of host plants. For more details of BBWV, refer to Vicia faba.

L

1386

Limonium spp. (Statice)

Carnation Italian ringspot virus Taxonomic position Genus: Tombusvirus

(CIRV)

Family: Tombusviridae

CIRV infection in plants of Limonium sinuatum was reported from Italy (Russo et al. 2008). The virus-infected statice plants exhibit symptoms of stunting, chlorosis, and distortion of young leaves that turned yellow or reddish with age. Some plants died before flowering. The virus is mechanically sap-transmissible. The virus is graft-transmissible, and the use of infected propagation material helps spread this virus. No insect vector is known for this virus. For more details of CIRV, refer to Dianthus caryophyllus.

Clover yellow vein virus Taxonomic position Genus: Potyvirus

(ClYVV)

Family: Potyviridae

ClYVV infection in plants of Limonium spp. was reported from Japan and from the USA in plants derived from Colombia (Inoue et al. 1985; Lawson et al. 1985a, b). The virus-infected statice plants showed vein yellowing, leaf malformation, and stunting. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ClYVV, refer to Trifolium spp.

Cucumber mosaic virus

(CMV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV infection in plants of Limonium spp. was reported from Southern Germany (Hein et al. 1976; Iino and Inoue 1984; Matsumoto et al. 1997). The virus-affected statice plants had leaves with more or less definite mosaic pattern on reduced growth. The virus is mechanically sap-transmissible to a large number of host plants, and a number of aphid species transmit this virus in a non-persistent manner. For more details of CMV, refer to Cucumis sativus.

Cymbidium ringspot virus Taxonomic position Genus: Tombusvirus

(CymRSV)

Family: Tombusviridae

CymRSV infection in plants of Limonium spp. was reported from Colombia (Koenig et al. 2004). The virus-infected statice plants show chlorotic lesions and dwarfing. The virus is transmitted in water and soil with no apparent vector. The virus is transmissible by mechanical sap-inoculation. For more details of CymRSV, refer to Cymbidium spp.

Limonium spp. (Statice)

1387

Grapevine Algerian latent virus Taxonomic position Genus: Tombusvirus

(GALV)

Family: Tombusviridae

GALV infection in plants of Limonium spp. was reported from the Netherlands and Japan (Koenig et al. 2004; Fujinaga et al. 2009). The virus-infected statice plants exhibit symptoms of dwarfing, leaf deformation, chlorotic leaf spot or necrosis, and flower distortion. There is no known vector for this virus. The virus is mechanically sap-transmissible. The primary spread of the virus takes place through the use of infected planting material. For more details of GALV, refer to Vitis vinifera.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Limonium sinuatum was reported from Italy (Vaira et al. 1992a, b, 1993). The virus-infected statice plants exhibit symptoms of necrosis, malformation, formation and yellow rings, flower or leaf break, yellowing on the foliage, and stunting of the plants; no flowers were produced. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Limonium flower distortion virus Taxonomic position Genus: Tombusvirus

(LFDV)

Family: Tombusviridae

Geographical distribution LFDV infection in plants of Limonium sinuatum was reported from the Netherlands (Koenig et al. 2004; Verhoeven et al. 2006). Symptoms and host(s) The virus-infected statice plants exhibit symptoms of dwarfing, leaf deformation, chlorotic leaf spot or necrosis, and flower distortion. Transmission The virus is transmitted in water and soil with no apparent vector. The virus is mechanically sap-transmissible. Virion properties and genome The virions are isometric and icosahedral (T = 3), non-enveloped, and about 32–35 nm in diameter. The genome is a monopartite, linear, single-stranded, positive-sense RNA. A partial genome sequence of 1231 nt is available (AY500882) (Verhoeven et al. 2006; White 2011).

L

1388

Limonium spp. (Statice)

Petunia asteroid mosaic virus Taxonomic position Genus: Tombusvirus

(PetAMV)

Family: Tombusviridae

PetAMV infection in plants of Limonium spp. was reported from the Netherlands (Koenig et al. 2004). The virus-infected statice plants cause chlorotic lesions, dwarfing, and necrosis. The virus is transmitted in water and soil with no apparent vector. The virus is mechanically sap-transmissible and also by grafting. For more details of PetAMV, refer to Petunia spp.

Sitke waterborne virus Taxonomic position Genus: Tombusvirus

(SWBV)

Family: Tombusviridae

Geographical distribution SWBV infection in plants of Limonium sinense was reported from Peru and Japan (Koenig et al. 2004; Horita et al. 2010). Symptoms and host(s) The virus-infected statice plants exhibit chlorotic lesions, leaf deformation and reddening, dwarfing, and tip necrosis symptoms. Transmission The virus is transmitted in water and soil with no apparent vector. The virus is mechanically sap-transmissible. Virion properties and genome The virions are isometric and icosahedral (T = 3), non-enveloped, and about 32–35 nm in diameter. The genome is a monopartite, linear, single-stranded, positive-sense RNA. A partial sequence of a limonium isolate is available (AY500886; 1194 nt) and a slightly longer sequence of another isolate (AJ607404; 1276 nt) (White 2011).

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in plants of Limonium spp. was reported from the Netherlands (Dijkstra and Van Dijke 1981). The virus-infected statice plants exhibited bright yellow or red line patterns and ring-spots on the leaves. The virus is transmitted by nematode vectors and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Limonium spp. (Statice)

1389

Tomato bushy stunt virus Taxonomic position Genus: Tombusvirus

(TBSV)

Family: Tombusviridae

TBSV infection in plants of Limonium spp. was reported from Germany (Krczal and Beutel 1994), Italy (Galetzka et al. 2000), and the Netherlands (Koenig et al. 2004). The virus-infected statice plants show mosaic, yellowing, necrosis, and malformation of leaves, together with severe flower damage. Severely infected plants ceased producing flowers or died. The virus is transmitted through soil but with no known vector. The virus is transmissible by mechanical sap-inoculation, and also by grafting. The virus is not transmissible by contact between plants. For more details of TBSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV has been reported to infecting plants of Limonium perezii in Italy (Vaira et al. 1993), from L. sinuatum in Spain (Jorda et al. 1995) and Portugal (Louro 1996), and L. latifolium in Lithuania (Samuitiene and Navalinskiene 2010). The virus-infected L. perezii plants showed diffuse yellowing, vein netting, and necrosis on the major veins of mature leaves. L. sinuatum showed leaf curling, stem and leaf necrosis, and general poor performance; no flowers were produced. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Limonium perezii plants was reported from the USA, Germany, and Japan (Niblett et al. 1969; Laird and Dickson 1972; Hein et al. 1976; Iino and Inoue 1984). The virus-infected statice plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

References Dijkstra J, van Dijke HD (1981) Line pattern in Limonium latifolium caused by Tobacco rattle virus. Neth J Plant Pathol 87:35–44 Fujinaga M, Ogiso H, Wakabayashi H, Morikawa T, Natsuaki T (2009) First report of a Grapevine Algerian latent virus disease on statice plants (Limonium sinuatum) in Japan. J Gen Plant Pathol 75:157–159 Galetzka D, Russo M, Rubino L, Krczal G (2000) Molecular characterization of a tombusvirus associated with a disease of statice (Goniolimon tataricum (L.) Boiss.). J Plant Pathol 82:151–155

L

1390

Linaria canadensis (Canadian toadflax)

Hein A, Koenig DE, Querfurth G (1976) Kohlschwarzringfleckigkeits-Virus und Gerkenmosaik-Virus in StaticeKulturen in Sȕddeutschland. Z Pflanzenkrankh, Pflanzenschutz 83:229–233 Hein A, Lesemann DE, Querfuth G (1977) Das Broad bean wilt-Virus in Limonium-Kulturen un Süddeutschland. J Phytopathol 89:340–346 Horita H, Matsui R, Sasaki J (2010) Sikte waterborne virus isolated from statice (Limonium sinense hybrids). Ann Phytopathol Soc Jpn 76:76 Iino N, Inoue N (1984) Characterization of Cucumber mosaic virus and Turnip mosaic virus isolated from statice (Limonium spp.). Ann Phytopathol Soc Jpn 50:116–117 Inoue N, Maeda T, Iino N, Mitsuhata K (1985) Characterization of Clover yellow vein virus isolated from statice (Limonium sinuatum) (abstract in Japanese). Ann Phytopathol Soc Jpn 51:354–355 Jorda C, Ortega A, Juarez M (1995) New hosts of tomato spotted wilt virus. Plant Dis 79:538 Koenig R, Verhoeven JTJ, Fribourg CE, Pfeilstetter E, Lesemann DE (2004) Evaluation of various species demarcation criteria in attempts to classify ten new tombusvirus isolates. Arch Virol 149:1733–1744 Krczal G, Beutel M (1994) A new soilborne virus disease in statice (Gonolium tartaricum). Acta Hortic 377:115–122 Laird EF, Dickson RC (1972) Turnip mosaic virus-vector relationships in field-grown statice, Limonium perezii. Plant Dis Reptr 56:722–725 Lawson RH, Brannigan MD, Foster J (1985a) A strain of Clover yellow vein virus infecting statice (Limonium sinuatum). Acta Hortic 164:233–240 Lawson RH, Brannigan MD, Foster J (1985b) Clover yellow vein virus in Limonium sinuatum. Phytopathology 75:899–906 Louro D (1996) Detection and identification of tomato spotted wilt virus and Impatiens necrotic spot virus in Portugal. Acta Hortic 431:99–105 Matsumoto J, Okamura N, Ohki ST (1997) Cucumber mosaic and broad bean wilt viruses isolated from Limonium sinuatum and Rudbeckia  hybrida. Ann Phytopathol Soc Jpn 63:13–15 Niblett CL, Paulus AO, Semancik JS (1969) A mosaic disease of statice caused by Turnip mosaic virus. Phytopathology 59:1166–1167 Russo M, De Stradis A, Vovlas C (2008) A disease of Limonium sinuatum caused by Carnation Italian ringspot virus. J Plant Pathol(00077NN) 90:400 Samuitiene M, Navalinskiene M (2010) The agents of viral diseases affecting Limonium Mill. plants. Meistų želdynų formavimas 1:145–150 Vaira AM, Lisa V, Dellavelle G (1992a) Distribution of tospovirus in Liguria, Italy. 5th International Plant virus epidemiology symposium. Viruses, Vectors and the Environment. Valenzano Bari, pp 23–24 Vaira AM, Gallo S, Lisa V (1992b) Nuove infezioni da due Tospovirus (tomato spotted wilt e Impatiens necrotic spot) in Liguria. Informatore Fitopatologco 42:37–42 Vaira AM, Roggero P, Luisoni E, Masenga V, Milne RG, Lisa V (1993) Characterization of two tospoviruses in Italy: tomato spotted wilt and impatiens necrotic spot. Plant Pathol 42:530–542 Verhoeven JTJ, Roenhorst JW, Lesemann DE, Koenig R (2006) Identification of tombusviruses isolated from Limonium sinuatum on the basis of host range, serological and molecular studies. Acta Hortic 722:201–208 White KA (2011) Tombusvirus. Tombusviridae. In: The Springer Index of Viruses. Springer, New York, pp 1917–1926. https://doi.org/10.1007/978-0-387-95919-1_314

Linaria canadensis (Canadian toadflax) Family: Scrophilariaceae

Weed host

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Linaria canadensis was reported from the USA (Groves et al. 2002). The virus-infected Canadian toadflax plants exhibit yellow spots or ring-spots on the leaves. The virus is

Lindernia anagallis

1391

transmitted by a thrips vector, Frankliniella fusca, in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Groves RL, Walgenbach JF, Mayor JW, Kennedy GG (2002) The role of weed hosts and tobacco thrips, Frankliniella fusca, in the epidemiology of Tomato spotted wilt virus. Plant Dis 86:573–582

Lindernia anagallis Family: Linderniaceae

Medicinal

Lindernia anagallis yellow vein virus Taxonomic position Genus: Begomovirus

(LaYVV)

L Family: Geminiviridae

Geographical distribution LaYVV infection in plants of Lindernia anagallis was reported from China and Vietnam (Che et al. 2007; Ha et al. 2008). Symptoms and host(s) The virus-infected lindernia plants exhibit severe mosaic, mottling, and blistering of leaves. Transmission The transmission of LaYVV has not been investigated. It is likely that, in common with other begomoviruses, LaYVV is transmitted by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The structure of the virions of LaYVV has not been investigated. In common with all geminiviruses, the virions of LaYVV are likely geminate (twinned quasi-icosahedra). LaYVV is a typical Old World monopartite begomovirus. The genome consists of a single molecule of a circular, single-stranded DNA (DNA-A) of 2740 nt (DQ641701 = NC_009550; AY795900, KC172827). A betasatellite of 1346 nt was found associated with the virus (DQ641715) has also been reported (Briddon 2001; Ha et al. 2008; Brown et al. 2015; Zerbini et al. 2017). The characterized genomes of LaYVV isolates encode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated.

1392

Linum usitatissimum (Flax)

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer Index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Che H-Y, Liu X-B, Yang X-G, Fu R-Y, Luo D-Q (2007) Molecular characterization of a distinct Begomovirus infecting Lindernia anagallis in China. Acta Phytopathol Sin 37:578–583 Ha C, Coombs S, Revill P, Harding R, Vu M, Dale J (2008) Molecular characterization of begomoviruses and DNA satellites from Vietnam: additional evidence that the New World geminiviruses were present in the Old World prior to continental separation. J Gen Virol 89:312–326 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Linum usitatissimum (Flax) Family: Linaceae

Oat blue dwarf virus Taxonomic position Genus: Marafivirus

Fiber crop

(OBDV)

Family: Tymoviridae

OBDV infection in plants of Linum usitatissimum was reported from Canada and the north central United States (Fredriksen and Goth 1959; Banttari and Moore 1962). The virus-infected flax plants exhibit the only leaves that are affected and display symptoms. A “crinkle” of diseased leaves occurs due to a swelling of lateral veins on the margins of the leaves, small indentations along the upper surfaces of the veins, and enations or pimples on the lower leaf surfaces. Diseased plants are also stunted and have reduced boll development and seed set. The virus is transmitted by the aster leafhopper vectors, Macrosteles fascifrons and M. laevis, in a persistent, propagative manner. The virus is not sap or seed-transmissible. For more details of OBDV, refer to Avena sativa.

Tobacco rattle virus Taxonomic position Genus: Tobravirus

(TRV)

Family: Virgaviridae

TRV infection in plants of Linum usitatissimum was reported from Egypt (Zein et al. 2012). The virusinfected flax plants exhibit yellowing, systemic mosaic, and leaf deformation. The virus is transmitted by nematode vectors, Paratrichodorus allius, P. anemones, P. christiei, P. nanus, P. pachydermus, P. teres, Trichodorus minor, and T. primitives. The virus is seed-transmitted in Linum usitatissimum up to 2.8–19.7% (Zein et al. 2012). The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Liriope spicata (Lilyturf)

1393

References Banttari EE, Moore MB (1962) Virus cause of blue dwarf of oats and its transmission to barley and flax. Phytopathology 52:897–902 Fredriksen RA, Goth RW (1959) Crinkle a new virus disease of flax. Phytopathology 49:538 Zein SN, Hamed AH, Zawam HS (2012) Tobacco rattle tobravirus: occurrence in flax plants (Linum usitatissimum L.) in Egypt. Nat Sci 10:14–20

Liparis bootanensis Family: Orchidaceae

Ornamental

Cymbidium mosaic virus Taxonomic position Genus: Potexvirus

(CymMV)

Family: Alphaflexiviridae

CymMV infection in plants of Liparis bootanensis was reported from India (Sherpa et al. 2003). The virus-infected liparis plants exhibit symptoms of flower color break, leaf necrosis, and necrotic spotting. The virus is mechanically sap-transmissible. The virus is also transmissible by contact between plants. No vector transmission is reported. The primary spread of the virus takes place through the use of virus-infected planting material. For more details of CymMV, refer to Cymbidium spp.

References Sherpa AR, Hallan V, Ram R, Vij SP, Pathak P, Garg ID, Zaidi AA (2003) First report of Cymbidium mosaic virus (CymMV) in orchids from India. Plant Pathol 52:788–788

Liriope spicata (Lilyturf) Family: Asparagaceae

Orchid fleck dichorhavirus Taxonomic position Genus: Dichorhavirus

Ornamental

(OFV)

Family: Rhabdoviridae

OFV infection in plants of Liriope spicata was reported from Australia (Mei et al. 2016). The virusinfected lilyturf plants exhibit symptoms of bright yellow flecks on the leaves. The virus is transmitted by mite vectors in a persistent, propagative manner, and also by mechanical sap-inoculation. For more details of OFV, refer to Cymbidium spp.

L

1394

Lithospermum arvense (Field gromwell)

References Mei MY, Bejerman N, Crew KS, McCaffrey N, Dietzgen RG (2016) First report of orchid fleck virus in lilyturf (Liriope spicata) in Australia. Plant Dis 100:1028

Lithospermum arvense (Field gromwell) Synonyms Buglossoides arvensis Family: Boraginaceae

Plum pox virus Taxonomic position Genus: Potyvirus

Weed host

(PPV)

Family: Potyviridae

PPV infection in plants of Lithospermum arvense was reported from Bulgaria (Milusheva and Rankova 2002). The virus-infected field gromwell plants exhibit pale or yellow green ring-spots or mottling on leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PPV, refer to Prunus domestica.

References Milusheva S, Rankova Z (2002) Plum pox potyvirus detection in weed species under field conditions. Acta Hortic 577:283–287

Litsea spp. Family: Lauraceae

Trees and shrubs

Bhendi yellow vein mosaic virus Taxonomic position Genus: Begomovirus

(BYVMV)

Family: Geminiviridae

BYVMV infection in plants of Litsea spp. was reported from India (Roy et al. 2015). The virus-infected litsea plants exhibit leaf curling, deformation, leaf yellowing, vein-clearing, mosaic, and stunted growth. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner.

Lobelia spp.

1395

The virus is not transmissible by mechanical inoculation. For more details of BYVMV, refer to Abelmoschus esculentus.

References Roy B, Chakraborty B, Mitra A, Sultana S, Sherpa AR (2015) Natural occurrence of Bhendi yellow vein mosaic virus on Litsea spp. in India. New Dis Rep 31:7

Lobelia spp. Family: Campanulaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Lobelia spp. was reported from the USA, France, Italy, and New Zealand (Nameth and Fisher 2001; Lockhart et al. 2002; Cardin et al. 2005). The virus-infected lobelia plants exhibit mosaic, leaf deformation, and yellow ring-spot symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV has been reported from plants of Lobelia spp. in the USA, Italy, and Finland (DeAngelis et al. 1993; Gotta et al. 1996; Lemmetty et al. 2011). The virus-infected lobelia plants exhibit symptoms of chlorotic flecking, mottle, chlorotic or necrotic spots and ring-spots, distortion, and terminal necrosis. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV infection in plants of Lobelia spp. was reported from the USA (Dobhal et al. 2015). The virus-infected lobelia plants may show either no significant symptoms or mottle-mosaic. There is no known vector for this virus. The virus is mechanically sap-transmissible and also by contact between plants. For more details of TMV, refer to Nicotiana tabacum.

L

1396

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

Lobelia spp.

(TRSV)

Family: Secoviridae

TRSV has been reported from plants of Lobelia spp. in the Netherlands (Lockhart et al. 2002; Sirca et al. 2007). The virus-infected lobelia plants exhibit symptoms of stunting, mosaic, necrosis, and deformation of leaves and color break of flowers. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Lobelia spp. was reported from Belgium, the Netherlands, and the USA (Gofflot and Verhoyen 1990; Verhoeven and Roenhorst 1994; Dobhal et al. 2015). The virus-infected lobelia plants exhibit symptoms of chlorotic flecks on leaves. The virus is transmitted by thrips vectors in a persistentpropagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Lobelia spp. was reported from commercial nurseries in the USA and New Zealand (Lockhart et al. 2002; Ochoa Corona et al. 2007). The virus-infected lobelia plants exhibit mosaic, leaf deformation, and mottle symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of TuMV, refer to Brassica rapa.

References Cardin L, Delecolle B, Moury B (2005) Occurrence of mosaic caused by Cucumber mosaic virus in lobelia hybrids in France and Italy. Plant Dis 89:340 DeAngelis JD, Sether DM, Rossignol PA (1993) Survival, development and reproduction in western flower thrips (Thysanoptera: Thripidae) exposed to Impatiens necrotic spot virus. Environ Entomol 22:1308–1312 Dobhal S, Arif M, Olson J, Mendoza-Yerbafría A, Aguilar-Moreno S, Perez-Garcia M, Ochoa-Corona FM (2015) Sensitive detection and discrimination method for studying multiple infections of five major plant viruses infecting ornamental plants in nursery environments. Ann Appl Biol 166:286–296 Gofflot A, Verhoyen M (1990) Speedy development of tomato spotted wilt virus infection transmitted by Frankliniella occidentalis in ornamental greenhouse cultivars in Belgium. Parasitica 46:85–88 Gotta P, Gallo S, Moretti F, Roggero P, Dellavalle G, Lisa V, Conti M (1996) Impatiens necrotic spot tospovirus in ornamentals in Piedmont (North-Western Italy) [Sinningia, Begonia, Impatiens, Lobelia, Eustonia]. Informatore Fitopatologico

Lolium spp. (Ryegrass; Italian ryegrass)

1397

Lemmetty A, Laamanen J, Soukainen M (2011) Emerging virus and viroid pathogen species identified for the first time in horticultural plants in Finland in 1997–2010. Agric Food Sci 20:29–41 Lockhart BEL, Fetzer J, Westendrop J (2002) Previously unreported viral diseases of Aster, Heuchera, Lobelia, Pulmonaria and Physostegia in the USA. Acta Hortic 568:221–224 Nameth SGP, Fisher JR (2001) First report of a Cucumber mosaic virus-associated satellite RNA in Lobelia erinus. Aust Plant Dis 85:802 Ochoa Corona FM, Lebas BSM, Elliott DR, Tang JZ, Alexander BJR (2007) New host records and new host family range for Turnip mosaic virus in New Zealand. Austra Plant Dis Notes 2:177–130 Sirca S, Stare BG, Pleško IM, Marn MV, Urek G, Javornik B (2007) Xiphinema rivesi from Slovenia transmit Tobacco ringspot virus and Tomato ringspot virus to cucumber bait plants. Plant Dis 91:770–770 Verhoeven JTJ, Roenhorst JW (1994) Tomato spotted wilt virus: ecological aspects in ornamental crops in the Netherlands from 1989 up to 1991. Acta Hortic 377:175–182

Lolium spp. (Ryegrass; Italian ryegrass) Family: Poaceae

Forage crop

Barley yellow dwarf virus

(BYDV)

Taxonomic position BYDV is a tentative member of the genus Luteovirus and family Luteoviridae BYDV infection in plants of Lolium perenne was reported from New Zealand, Scotland, and Australia (Latch 1977; Guy et al. 1986; Eagling et al. 1989; Henry and Dedryver 1991; Holmes 1991; Henry et al. 1992; Dempster and Holmes 1995). The virus-infected ryegrass plants do not show obvious symptoms. The virus is transmitted by a number of aphid species in a circulative, non-propagative manner. No mechanical transmission of this virus is reported. For more details of BYDV, refer to Hordeum vulgare.

Barley yellow dwarf virus PAV (BYDV-PAV) Taxonomic position Genus: Luteovirus

Family: Luteoviridae

BYDV-PAV infection in plants of Lolium rigidum was reported from Syria (Ansi et al. 2007). The virus is transmitted by aphid vectors in a circulative non-propagative manner. The virus is not transmitted by mechanical inoculation. For more details of BYDV-PAV, refer to Hordeum vulgare.

Cereal yellow dwarf virus RPV Taxonomic position Genus: Polerovirus

(CYDV-RPV)

Family: Luteoviridae

CYDV-RPV infection in plants of Lolium perenne plants was reported from Latvia (Bisnieks et al. 2006). The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of CYDV-RPV, refer to Hordeum vulgare.

L

1398

Lolium latent virus Taxonomic position Genus: Lolavirus

Lolium spp. (Ryegrass; Italian ryegrass)

(LoLV)

Family: Alphaflexiviridae

Geographical distribution LoLV infection in plants of Lolium spp. was reported from Europe (Germany, the Netherlands, France, and the United Kingdom) and from the United States (Huth et al. 1995; Li et al. 2005; Maroon-Lango et al. 2006; Vaira et al. 2008). Symptoms and host(s) The virus-infected plants of ryegrass are generally symptomless but show faint chlorotic mottle/ flecking on leaves transiently in the spring when conditions are suitable. The natural host range of the virus is restricted to gramineaceous species including ryegrass (lolium). It also readily infects N. benthamiana and a few other dicotyledonous species. In N. benthamiana, it induces local lesions and a systemic mosaic that varies from mildly chlorotic to white. Transmission The virus is very inefficiently transmitted by the aphid vector Rhopalosiphum padi (Huth et al. 1995). The virus is readily transmissible to healthy host plants by sap-inoculation, and infects 25 plant species in 5 families tested. The virus is transmitted by vegetative propagation. No pollen and seed-transmission is reported. Virion properties and genome The virions are slightly flexuous filaments of 640 nm modal length and about 13 nm wide. The virions contain a single molecule of linear positive-sense ssRNA, 7674 nt (EU489641 = NC_010434), excluding the poly(A) tail (Vaira et al. 2008).

Ryegrass cryptic virus Taxonomic position Genus: Unassigned

(RGCV)

Family: Partitiviridae

Geographical distribution RGCV infection in plants of Lolium spp. was reported from Australia, England, Germany, Italy, and the United Kingdom (Plumb 1973; Luisoni and Milne 1980; Guy and Sward 1991). Symptoms and host(s) Symptoms of this disease, in ryegrass, include chlorotic spots accompanied by necrosis, particularly on the oldest leaves. The natural hosts of this virus are Lolium multiflorum and L. perenne. Transmission There is no known natural vector for this virus. The virus is not mechanically sap-transmissible and also not by grafting. The virus is transmitted by seed in L. multiflorum up to 82% (Plumb 1973; Plumb and Lennon 1980). The virus is pollen transmitted.

Lolium spp. (Ryegrass; Italian ryegrass)

1399

Virion properties and genome The virions are isometric and 29–30 nm in diameter. The genome typically consists of two dsRNA segments (Luisoni and Milne 1980; Vainio et al. 2018). No sequence information is available.

Ryegrass mosaic virus Taxonomic position Genus: Rymovirus

(RGMV)

Family: Potyviridae

Geographical distribution RGMV infection was first reported in plants of Lolium spp. from Washington State, USA (Bruehl et al. 1957). The virus is reported from the USA, Canada, Australia, England, France, Holland, Germany, Denmark, Sweden, Victoria, New Zealand, Lithuania, and Finland (Guy and Sward 1991; Eagling et al. 1992; Smales et al. 1996; Webster et al. 1999; Guy 2006; Urbanaviciene 2008). Symptoms and host(s) The virus-infected ryegrass plants exhibit pale green streaks and mosaic in the leaf lamina and stunting of plants, and in severe cases, the leaf may show yellow flecking which is followed by necrosis and death of the tillers. The host range of RGMV is restricted to festucoid species within the family Poaceae. Ryegrass species L. multiflorum and L. perenne are the most important hosts, but other grasses such as cocksfoot (Dactylis glomerata), fescue (Poa annua) and Agrostis stolonifera can be naturally infected. Transmission The virus is transmitted by the mite vector Abacarus hystrix which acquires the virus from infected plants in 2 h and loses the ability to transmit the virus within 24 h of leaving the infected plant (Mulligan 1960; Lewis and Heard 1980; Gibson 1981). Adult mites are wind-borne from infected onto healthy plants, the primary means of RGMV spread (Gibson 1980). The virus is transmissible by mechanical inoculation, but there is no evidence of seed and pollen transmission. This virus is also transmitted through farm machinery and livestock (Catherall 1987). The perennial Lolium perenne is particularly important for the reservation of infectivity and transmission of RGMV into the next cropping season. Virion properties and genome The virions are non-enveloped, flexuous filaments 700 nm long and 15 nm in diameter. The genome consists of a single molecule of linear, positive-sense, single-stranded RNA of 9535 nt (Y09854 = NC_001814) with a 30 -poly(A) terminus (Zagula et al. 1992; Schubert et al. 1995, 1999, 2005; Lopez-Moya et al. 2009; Wylie et al. 2017).

Ryegrass mottle virus

(RGMoV)

Synonyms Lolium mottle virus Taxonomic position Genus: Sobemovirus

Family: Solemoviridae

L

1400

Lolium spp. (Ryegrass; Italian ryegrass)

Geographical distribution RGMoV-infected plants of Lolium spp. were reported for the first time from Japan by (Toriyama et al. 1983). The virus is present in Western Europe and Germany (Zhang et al. 2001). Symptoms and host(s) The virus-infected ryegrass plants exhibit symptoms first at the base of younger leaves as mild chlorotic mottling, which later encompasses the whole plant. The symptom in mature plants is longitudinal necrosis on older leaves. The natural hosts of this virus are Lolium perenne and Cynosurus cristatus. Other susceptible plants include Avena sativa, Hordeum vulgare, Lagurus ovatus, Lolium multiflorum, and Phleum pratense. Transmission There is no known vector for this virus. The virus is mechanically sap-transmissible to certain members of Gramineae family. Virion properties and genome The virions are isometric, non-enveloped, and 28 nm in diameter. The genome is a polycistronic, positive-sense, single-stranded RNA (ssRNA) genome of 4210 nt (EF091714 = NC_003747; AB040446). The genome consists of two overlapping ORFs, ORF2a and ORF2b (Zhang et al. 2001; Balke et al. 2007; Plevka et al. 2007; Somera et al. 2015).

References Ansi A, Kumari SG, Haj Kasem A, Makkouk KM, Muharram I (2007) The occurrence of barley yellow dwarf viruses on cereal crops and wild grasses in Syria. Arab J Pl Prot 25:1–9 Balke I, Resevica G, Zeltins A (2007) The ryegrass mottle virus genome codes for a sobemovirus 3C-like serine protease and RNA-dependent RNA polymerase translated via -1 ribosomal frameshifting. Virus Genes 35(2):395–398 Bisnieks M, Kvarnheden A, Turka I, Sigvald R (2006) Occurrence of barley yellow dwarf virus and cereal yellow dwarf virus in pasture grasses and spring cereals in Latvia. Acta Agric Scand Sect B Soil Plant Sci 56:171–178 Bruehl GW, Toko H, McKinney HH (1957) Mosaics of Italian ryegrass and orchard grass in western Washington. Phytopathology 47:517 Catherall PL (1987) Effects of barley yellow dwarf and ryegrass mosaic viruses alone and in combination on the productivity of perennial and Italian ryegrass. Plant Pathol 36:73–78 Dempster LC, Holmes SJI (1995) The incidence of strains of barley yellow dwarf virus in perennial ryegrass crops in south west and central Scotland. Plant Pathol 44:710–717 Eagling DR, Cunningham PJ, Sward RJ, Halloran GM (1989) The incidence of barley yellow dwarf virus isolates in perennial ryegrass (Lolium perenne) in south-western Victoria. Plant Pathol 38:408–413 Eagling DR, Villalta O, Sward RJ (1992) Host range, symptoms and effects on pasture production of a Victorian isolate of ryegrass mosaic potyvirus. Aust J Agric Res 43(5):1243–1251 Gibson RW (1980) Interactions between methods of cultivating ryegrass and the spread of Ryegrass mosaic virus. 3rd Conf Virus Dis Gramineae Europe, Rothamsted, 28–30 May Gibson RW (1981) Rapid spread by mites of ryegrass mosaic virus from old sward to seedlings of ryegrass and its prevention by aldicarb. Plant Pathol 30:25–29 Guy PL (2006) New Zealand grasslands revisited: identification of cocksfoot mild mosaic virus. Australas Plant Pathol 35:461–464 Guy PL, Sward RJ (1991) Ryegrass mosaic and ryegrass cryptic in Australia. Acta Phytopathol Entomol Hung 26:199–202 Guy PL, Johnstone GR, Duffus JE (1986) Occurrence and identity of barley yellow dwarf viruses in Tasmanian pasture grasses. Aust J Agric Res 37:43–53 Henry M, Dedryver CA (1991) Occurrence of barley yellow dwarf virus in pastures of western France. Plant Pathol 40:93–99 Henry M, Francki RIB, Wallwork H (1992) Occurrence of barley yellow dwarf virus in cereals and grasses of the lowrainfall wheat belt of South Australia. Plant Pathol 41(6):713–721

Lonicera spp. (Honeysuckle)

1401

Holmes SJI (1991) Barley yellow dwarf virus in Lolium spp. Acta Phytopathol Entomol Hung 26:33–39 Huth W, Lesemann DE, Gotz R, Vetten HJ (1995) Some properties of Lolium latent virus. Agronomie 15:508 Latch GCM (1977) Incidence of barley yellow dwarf virus in ryegrass pastures in New Zealand. N Z J Agric Res 20:87–89 Lewis CG, Heard JA (1980) The incidence of ryegrass crops of eriophyid mites, vectors of ryegrass mosaic virus. 3rd Conf Virus Dis Gramineae Europe, Rothamsted, 28–30 May Li R, Mock R, Maroon-Lango C, Hammond J (2005) Partial characterization of a strain of Lolium latent virus, a Flexivirus from ryegrass (Lolium perenne). Second Asian Conference of Plant Pathology, Singapore, June 2005, p 89 Lopez-Moya JJ, Valli A, Garcia JA (2009) Potyviridae. In: Encyclopaedia of Life Sciences (ELS). Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000755.pub2 Luisoni E, Milne RG (1980) Some properties of ryegrass spherical virus. Proc 3rd conf on virus diseases of Gramineae in Europe. Rothamsted Exp Stn., Harpenden, Herts, UK, p 73 Maroon-Lango CJ, Hammond J, Warnke S, Li R, Mock R (2006) First report of Lolium latent virus in ryegrass in the United States. Plant Dis 90:528 Mulligan TE (1960) The transmission by mites, host-range and properties of ryegrass mosaic virus. Ann Appl Biol 48:575–579 Plevka P, Tars K, Zeltins A, Balke I, Truve E, Liljas L (2007) The three-dimensional structure of ryegrass mottle virus at 2.9 a resolution. Virology 369(2):364–374 Plumb RT (1973) Ryegrass spherical virus. Rothamsted Exp Station Rep 1972 1:125 Plumb RT, Lennon EA (1980) Seed transmission and effects of ryegrass spherical virus. Proc 3rd conf on virus diseases of Gramineae in Europe. Rothamsted Exp Stn., Harpenden, Herts, UK, pp 75–79 Schubert J, Rabenstein F, Proll E (1995) Sequence of the 30 -part of the RNA of ryegrass mosaic virus, a potyvirus. Agronomie 15:447–452 Schubert J, Fauquet C, Merits A, Rabenstein F (1999) The complete nucleotide sequence of the ryegrass mosaic potyvirus indicates that it is a recombinant between members of two different genera in the family Potyviridae. J Plant Dis Protect 106(4):392–404 Schubert J, Merits A, Jaervekuelg L, Paulin L, Rubenstein F (2005) The complete nucleotide sequence of the ryegrass mosaic virus genomic RNA. DDBJ/EMBL/GenBank databases. Accession number (Y09854). Smales TE, Ferguson CM, Guy PL, Shand JA (1996) A survey of ryegrass mosaic virus and endophyte in Otago and Southland. In: Proceedings of the 49th New Zealand Plant Protection Conference. pp 220–224 Somera M, Sarmiento C, Truve E (2015) Overview on Sobemoviruses and a proposal for the creation of the family Sobemoviridae. Viruses 7:3076–3115 Toriyama S, Mikoshiba Y, Doi Y (1983) Ryegrass mottle virus, a new virus from Lolium multiflorum in Japan. Ann Phytopathol Soc Jpn 49:610–618 Urbanaviciene L (2008) Identification of ryegrass mosaic rymovirus in poaceae plants. Biologia 54:75–78 Vainio EJ, Chiba S, Ghabrial SA, Maiss E, Roossinck M, Sabanadzovic S, Suzuki N, Xie J, Nibert M, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Partitiviridae. J Gen Virol 99:17–18 Vaira AM, Maroon-Lango CJ, Hammond J (2008) Molecular characterization of Lolium latent virus, proposed type member of a new genus in the family Flexiviridae. Arch Virol 153:1263–1270 Webster DE, Guy PL, Beck DL, Forster RLS (1999) Distribution and diversity of New Zealand isolates of ryegrass mosaic virus. Arch Virol 144:2059–2064 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Zagula KR, Niblett CL, Robertson NL, French R, Lommen SA (1992) Potyviridae: genus Rymovirus. Arch Virol 5:269–276 Zhang F, Toriyama S, Takahashi M (2001) Complete nucleotide sequence of ryegrass mottle virus: a new species of the genus Sobemovirus. J Gen Plant Pathol 67:63–68

Lonicera spp. (Honeysuckle) Family: Caprifoliaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

L

1402

Lonicera spp. (Honeysuckle)

CMV infection in plants of Lonicera spp. was reported from the United Kingdom, the Netherlands, Poland, and Sweden (Lihnell 1951; Kaminska et al. 2005). The virus-infected honeysuckle plants show severe leaf chlorosis, stunted growth, distorted leaves and flowers, and ring-spots. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Eggplant mottled dwarf nucleorhabdovirus Taxonomic position Genus: Nucleorhabdovirus

(EMDV)

Family: Rhabdoviridae

EMDV infection in plants of plants of Lonicera spp. was reported from Tunisia (Martelli et al. 1987). The virus-infected honeysuckle plants exhibit vein yellowing symptoms. The virus is mechanically sap-transmissible. The virus is transmitted by leafhopper vectors in a persistent, propagative manner. The virus is mechanically sap-transmissible and also by grafting. For more details of EMDV, refer to Solanum melongena.

Honeysuckle ringspot virus Taxonomic position Genus: Alphacarmovirus

(HnRSV)

Family: Tombusviridae

Geographical distribution HnRSV infection in plants of Lonicera spp. was reported from California, USA (Gulati-Sakhuja et al. 2011). Symptoms and host(s) The virus induces yellow to purple rings in infected honeysuckle plants. Transmission The virus is mechanically sap-transmissible to Chenopodium murale. Virion properties and genome The virions are isometric and 29–31 nm in diameter and have T = 3 icosahedral symmetry. The genome consists of a single molecule of linear, positive-sense ssRNA of 3956 nt (HQ677625 = NC_014967) (Gulati-Sakhuja et al. 2011).

Honeysuckle yellow vein virus

(HYVV)

Synonyms Honeysuckle yellow vein Kagoshima virus (HYVKgV) Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Lonicera spp. (Honeysuckle)

1403

Geographical distribution HYVV infection in plants of Lonicera japonica was reported from Japan, Korea, the United Kingdom, and New Zealand (Briddon 2002; Lyttle and Guy 2004; Ueda et al. 2008; Wang et al. 2011). Symptoms and host(s) The virus-infected honeysuckle plants exhibit bright yellow vein symptoms (yellowing of main and small veins) which develop into a bright foliar yellow mosaic. HYVV has been identified in Lonicera japonica (var. aureoreticulata), tomato (Solanum lycopersicum), and tobacco (Nicotiana tabacum). Transmission The transmission of HYVV has not been investigated. It is likely that, in common with other begomoviruses, HYVV is transmitted by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner. The virus is transmitted by cuttings and by grafting. The virus is not transmissible by contact between plants. Isolates of the virus in Europe, and possibly elsewhere, are not transmissible by B. tabaci, which is likely due to the long maintenance of Lonicera japonica (var. aureoreticulata) by vegetative means, in the absence of the whitefly vector (Bedford et al. 1994). Experimentally, the virus has been shown to infect Nicotiana benthamiana by Agrobacterium-mediated inoculation of the clone viral genome (Lee et al. 2011). Virion properties and genome The structure of the virions of HYVV has not been investigated. In common with all geminiviruses, the virions of HYVV are likely geminate (twinned quasi-icosahedra). HYVV is a typical Old World monopartite begomovirus. The genome consists of a single circular molecule of a single-stranded DNA of 2784 nt (AJ542540 = NC_005807; AB182261) (Briddon 2001; Wang et al. 2011; Brown et al. 2015; Zerbini et al. 2017). Honeysuckle yellow vein betasatellite DNA molecule is associated with HYVV, and consists of 1344 nt (AJ316040) (Briddon et al. 2003; Zhou 2013). The characterized genomes encode the six genes typically encoded by monopartite begomoviruses originating from the Old World. The expression and function of the genes have not been investigated. HYVV associates with a betasatellite (Ogawa et al. 2008; Saunders et al. 2008).

Prune dwarf virus Taxonomic position Genus: Ilarvirus

(PDV)

Family: Bromoviridae

PDV infection in plants of Lonicera caucasica, L. maackii, and L. sachalinensis was reported from Hungary (Nemeth et al. 2010). The virus-infected honeysuckle plants show symptoms of pale green spots and vein-clearing. The virus is transmitted by the thrips vectors; the virus is present in/on pollen and entering the host through injuries caused by thrips while feeding. The virus is transmissible by mechanical sap-inoculation. For more details of PDV, refer to Prunus avium.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

L

1404

Lonicera spp. (Honeysuckle)

TSV was reported from a plant of Lonicera fragrantissima grown in the USA from seed imported from China in 1914 (Waterworth 1998). The virus-infected honeysuckle plants showed chlorosis on some shoot tips and mild veinal chlorosis on other leaves. The virus is transmitted by the thrips vectors, and they assist in the transfer of virus-carrying pollen to the healthy plants through feeding wounds. The virus is mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato leaf curl Japan virus

(ToLCJV)

Synonyms Honeysuckle yellow vein mosaic virus (HYVMV); Tomato yellow dwarf virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

ToLCJV/HYVMV infection in plants of Lonicera spp. was reported from Japan, Korea, Europe, and Australia (Kitamura et al. 2004). The virus-infected honeysuckle plants exhibit bright yellow vein symptoms (yellowing of main and small veins followed by mosaic pattern), leaf crinkling, and stunting symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, nonpropagative manner. For more details of ToLCJV, refer to Solanum lycopersicum.

References Ali A, Ahmed M, Nishigawa H, Natsuaki T (2014) Identification of Tobacco leaf curl virus affecting Lonicera japonica, an ornamental plant common in Japan. J Agric Sci Technol 16:645–655 Bedford ID, Briddon RW, Brown JK, Rosell RC, Markham PG (1994) Geminivirus transmission and biological characterisation of Bemisia tabaci (Gennadius) biotypes from different geographic regions. Ann Appl Biol 125(2):311–325 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer Index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Briddon RW (2002) Diversity of European begomoviruses: identification of a new disease complex. EPPO Bull 32(1):1–5 Briddon RW, Bull SE, Amin I, Idris AM, Mansoor S, Bedford ID, Dhawan P, Rishi N, Siwatch SS, Abdel-Salam AM, Brown JK, Zafar Y, Markham PG (2003) Diversity of DNA beta, a satellite molecule associated with some monopartite begomoviruses. Virology 312(1):106–121 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6): 1593–1619 Gulati-Sakhuja A, Rains L, Tian T, Liu H (2011) The complete nucleotide sequence and genome organization of a novel carmovirus – honeysuckle ringspot virus isolated from honeysuckle. Arch Virol 156:1635–1640 Kaminska M, Silwa H, Malinowski T (2005) Partial characterization of Cucumber mosaic virus isolate infecting Lonicera caprifolium L. plants. Acta Sci Pol Hortorum Cultus 4:3–10 Kitamura K, Murayama A, Ikegami M (2004) Evidence for recombination among isolates of Tobacco leaf curl Japan virus and Honeysuckle yellow vein mosaic virus. Arch Virol 149:1221–1229 Lee G, Kim S, Jung J, Auh C-K, Choi E, Chang M, Lee S (2011) Agroinoculation of Nicotiana benthamiana with cloned honeysuckle yellow vein virus isolated from Lonicera japonica. Arch Virol 156(5):785–791

Lophanthus anisatus (Giant hyssop)

1405

Lihnell D (1951) Some host plants of cucumber virus 1 in Sweden. Vaxtskyddsnotiser 4:52–56 Lyttle DJ, Guy PL (2004) First record of Geminiviruses in New Zealand: Abutilon mosaic virus and Honeysuckle yellow vein virus. Australas Plant Pathol 33:321–322 Martelli G, Cherif C, Lafortezza R, Ezzaier K (1987) Eggplant mottled dwarf virus associated with vein yellowing of honeysuckle. J Phytopathol Phytopathol-Zeirscherift 119:32–41 Nemeth M, Nyerges K, Hangyál R, Kósa G (2010) Surveying viruses on ornamental trees and shrubs in two Hungarian botanical gardens and an arboretum. Julius-Kühn-Archiv 427:293–299 Ogawa T, Sharma P, Ikegami M (2008) First report of a strain of Tobacco leaf curl Japan virus associated with a satellite DNA in honeysuckle in Japan. Plant Pathol 57:391 Osaki T, Kobatake H, Inouye T (1979) Yellow vein mosaic of honeysuckle (Lonicera japonica thumb), a disease caused by Tobacco leaf curl virus in Japan. Ann Phytopathol Soc Jpn 45:62–69 Saunders K, Briddon RW, Stanley J (2008) Replication promiscuity of DNA-b satellites associated with monopartite begomoviruses; deletion mutagenesis of the Ageratum yellow vein virus DNA-b satellite localises sequences involved in replication. J Gen Virol 89(12):3165–3172 Ueda S, Onuki M, Hanada K, Takanami Y (2008) Unique grouping of the Far East Asian begomovirus complex based on sequence analyses of the DNA-A genome and associated DNAb satellite molecules isolated from tomato, honeysuckle and Eupatorium plants in Japan. Arch Virol 153(3):417–426 Wang Y, Ji J, Oh T, Oh S, Kim S, Lee H, Shim M, Choi C, Kim S, Kim I, Kim Y (2011) Occurrence of Honeysuckle Yellow Vein Virus (HYVV) containing a monopartite DNA-A genome in Korea. Eur J Plant Pathol 129:361–370 Waterworth HE (1998) First report of tobacco streak Ilarvirus from honeysuckle. Plant Dis 82:1402 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zhou X (2013) Advances in understanding begomovirus satellites. Annu Rev Phytopathol 51:357–381

L Lophanthus anisatus (Giant hyssop) Family: Lamiaceae

Medicinal

Tomato spotted wilt orthotospovirus (TSWV) Taxonomic position Genus: Orthotospovirus

Family: Tospoviridae

TSWV infection in plants of Lophanthus anisatus was reported from Ukraine (Dikova et al. 2016). The virus-infected giant hyssop plants exhibit symptoms of yellow mosaic on the leaves. The virus is transmitted by thrips vectors in persistent-propagative manner and also by mechanical sap-inoculation to large number of herbaceous hosts. The virus is transmitted by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Dikova B, Mishchenko L, Dunish A, Dashchenko A (2016) Tomato spotted wilt virus on Giant hyssop and common Valerian in Ukraine and Bulgaria. Bulgarian J Agric Sci 22(1):108–113

1406

Lotus corniculatus (Bird’s-foot trefoil)

Lotus corniculatus (Bird’s-foot trefoil) Family: Fabaceae

Forage crop

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Lotus corniculatus was reported from Yugoslavia (Buturovic 1974). The virus-infected bird’s-foot trefoil plants exhibit symptoms of mosaic, mottle, leaf distortion, and stunting. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in plants of Lotus corniculatus was reported from the USA and Europe (Ostazeski 1965; Bode and Klinkowski 1968). The virus-infected bird’s-foot trefoil plants do not show symptoms. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sapinoculation. The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV infection in plants of Lotus corniculatus was reported from the USA (Ostazeski and Scott 1966). The virus-infected bird’s-foot trefoil plants exhibit symptoms of deformed leaves covered with angular necrotic areas. The virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ToRSV, refer to Solanum lycopersicum.

References Bode O, Klinkowski M (1968) Handelspflanzen. In: Klinkowski M et al (eds) Pflanzliche Virologie, vol II/I. AkademieVerlag, Berlin, pp 161–187 Buturovic D (1974) Neke karakteristike jednog virusnog izolata iz smiljkite. Mikrobiologia 11:173–180 Ostazeski SA (1965) The natural occurrence of tobacco ringspot virus in bird’s-foot trefoil (Lotus corniculatus). Plant Dis Reptr 49:855–856 Ostazeski SA, Scott HA (1966) Natural occurrence of Tomato ringspot virus in bird’s-foot trefoil. Phytopathology 56:585–586

Ludwigia spp. (Primrose-willow)

1407

Ludwigia spp. (Primrose-willow) Family: Onagraceae

Weed host

Ludwigia yellow vein Vietnam virus Taxonomic position Genus: Begomovirus

(LuYVVNV)

Family: Geminiviridae

Geographical distribution LuYVVNV infection in plants of Ludwigia hyssopifolia was reported from China and Vietnam (Huang et al. 2006; Ha et al. 2008). Symptoms and host(s) The virus-infected primrose-willow plants exhibit yellow vein symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate), incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular, single-stranded DNA (DNA-A) of 2751 nt (DQ641699 = NC_038451) (Ha et al. 2008; Zerbini et al. 2017).

Ludwigia yellow vein virus Taxonomic position Genus: Begomovirus

(LuYVV)

Family: Geminiviridae

Geographical distribution LuYVV infection in plants of Ludwigia hyssopifolia was reported from China and Vietnam (Huang et al. 2006; Ha et al. 2008). Symptoms and host(s) The virus-infected primrose-willow plants exhibit yellow vein symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular, single-stranded DNA (DNA-A) of 2758 nt (AJ965539 = NC_007210) (Briddon 2001; Ha et al. 2008; Brown et al. 2015; Zerbini et al. 2017).

L

1408

Luffa acutangula (Ridge gourd)

Rice necrosis mosaic virus Taxonomic position Genus: Bymovirus

(RNMV)

Family: Potyviridae

RNMV infection in plants of Ludwigia perennis was reported from India (Ghosh 1981). The virus is transmitted by a fungal vector in a persistent manner. The virus is also transmissible by mechanical sap-inoculation. For more details of RNMV, refer to Oryza sativa.

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer Index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Ghosh SK (1981) Weed hosts of rice necrosis mosaic virus. Plant Dis 65:602–603 Ha C, Coombs S, Revill P, Harding R, Vu M, Dale J (2008) Molecular characterization of begomoviruses and DNA satellites from Vietnam: additional evidence that the New World geminiviruses were present in the Old World prior to continental separation. J Gen Virol 89:312–326 Huang JF, Jiang T, Zhou XP (2006) Molecular characterization of begomoviruses infecting Ludwigia hyssopifolia. J Plant Pathol 88(1):83–88 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Luffa acutangula (Ridge gourd) Family: Cucurbitaceae

Vegetable

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

CGMMV infection in plants of Luffa acutangula was reported from India and Taiwan (Hseu et al. 1987; Sharma et al. 2014; Kumar et al. 2017). The virus-infected ridge gourd plants exhibit symptoms of systemic mosaic and mottling on leaves, blistering of fruit, and deterioration of fruit pulp. The virus is not transmitted by any insect vector. The virus is transmitted through soil and irrigation water contaminated with infected plant debris. For more details of CGMMV, refer to Cucumis sativus.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

Luffa acutangula (Ridge gourd)

1409

CMV infection in plants of Luffa acutangula occurs wherever the crop is grown (Goel and Varma 1973; Huang et al. 1987; Kiranmai et al. 1998). The virus-infected ridge gourd plants at the early stages of the disease usually show a mosaic and chlorotic or irregular mottling of leaves, but symptoms become milder or disappear later in the season. All species were susceptible although the symptoms were masked at the late stages of growth. The virus is transmitted by a number of aphid species in a nonpersistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Luffa yellow mosaic virus Taxonomic position Genus: Begomovirus

(LYMV)

Family: Geminiviridae

Geographical distribution LYMV infection in plants of Luffa acutangula was reported from southern Vietnam (Revill et al. 2003). Symptoms and host(s) The virus-infected ridge gourd plants exhibit yellow mosaic symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2742 nt (AF509739 = NC_004824) and DNA-B of 2713 nt (AF509740 = NC_004825) (Briddon 2001; Revill et al. 2003; Brown et al. 2015; Zerbini et al. 2017).

Papaya ringspot virus Taxonomic position Genus: Potyvirus

(PRSV)

Family: Potyviridae

PRSV infection in plants of Luffa acutangula was reported from Tamilnadu (India) (Nagendran et al. 2017). The virus is transmitted by aphid vectors in a non-persistent manner and also by mechanical sapinoculation. For more details of PRSV, refer to Carica papaya.

Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

ToLCNDV infection in plants of Luffa acutangula was reported from India (Tiwari et al. 2011; Manjunath 2014; Manjunath et al. 2016; Rajeshwari et al. 2016; Patil et al. 2017). The virus-infected ridge gourd plants exhibit mosaic mottling, leaf curl, and extensive chlorosis symptoms. The virus is

L

1410

Luffa acutangula (Ridge gourd)

transmitted by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner. For more details of ToLCNDV, refer to Solanum lycopersicum.

Watermelon bud necrosis orthotospovirus Taxonomic position Genus: Orthotospovirus

(WBNV)

Family: Tospoviridae

WBNV was identified in plants of Luffa acutangula showing yellowing of leaves in southern India (Mandal et al. 2003). The virus was mechanically transmissible to ridge gourd plants, which showed chlorotic spots on the inoculated leaves and vein-clearing followed by chlorotic rings and yellow netting along veins of the newly developed leaves. The virus is transmitted by thrips vectors in a persistent, propagative manner, and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of WBNV, refer to Citrullus lanatus.

Zucchini yellow mosaic virus Taxonomic position Genus: Potyvirus

(ZYMV)

Family: Potyviridae

ZYMV infection in plants of Luffa acutangula was reported from Pakistan and Taiwan (Hseu et al. 1987; Ashfaq et al. 2015). The virus-infected ridge gourd plants exhibit mosaic, leaf malformation, and blistering symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ZYMV, refer to Cucurbita pepo.

References Ashfaq M, Saeed U, Mukhtar T, ul Haq MI (2015) First report of Zucchini yellow mosaic virus in ridge gourd in Pakistan. Plant Dis 99:1870 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer Index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Goel RK, Varma JP (1973) Mosaic disease of ridge gourd (Luffa acutangula Roxb.) in Haryana. Haryana Agric Univ J Res 3:135–144 Hseu SH, Huang CH, Chang CA, Yang WZ, Chang YM, Hsiao CH (1987) The occurrence of five viruses in six cucurbits in Taiwan. Plant Protect Bull 29:233–244 Huang CH, Chao YJ, Chang CA, Hseu SH, Hsaio CH (1987) Identification and comparison of different viruses on symptom expression in Luffa. J Agric Res China 36:413–420 Kiranmai G, Sreenivasulu P, Nayudu MV (1998) Epidemiology of cucumber mosaic cucumovirus isolates naturally infecting three solanaceous vegetable crops around Tirupati. Ind Phytopathol 51:315–318 Kumar A, Jailani AAK, Roy A, Mandal B (2017) The occurrence, biology and genomic properties of tobamoviruses infecting crop plants in India. In: Mandal B, Rao GP, Baranwal VK, Jain RK (eds) A century of plant virology in India. Springer Nature, Singapore, pp 429–443. (ISBN 978-981-10-5671-0) Mandal B, Jain RK, Chaudhary V, Varma A (2003) First report of natural infection of Luffa acutangula by Watermelon bud necrosis virus in India. Plant Dis 87:598

Luffa cylindrica (Sponge gourd, Suakwa)

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Manjunath SH (2014) Biological and molecular characterization of yellow moaic virus infecting Ridge gourd (Luffa acutangula) and Cucumber (Cucumis sativus). MSc thesis submitted to the Department of Plant Pathology University of Agricultural Sciences, Bengaluru (India) Manjunath SH, Rangaswamy KT, Basavaraj S, Bhagyashree M, Nagaraju N, Prameela HA (2016) Molecular detection and identification of tomato leaf curl New Delhi virus associated with yellow mosaic disease of ridge gourd (Luffa accutangula L.) based on coat protein gene. Int J Agric Sci 8:3444–3449 Nagendran K, Mohankumar S, Aravintharaj R, Balaji CG, Manoranjitham SK, Singh AK, Rai AB, Singh B, Karthikeyan G (2017) The occurrence and distribution of major viruses infecting cucurbits in Tamil Nadu state, India. Crop Prot 99:10–16 Patil CV, Ramdas SV, Premchand U, Shankarappa KS (2017) Survey, symptomatology, transmission, host range and characterization of begomovirus associated with yellow mosaic disease of ridge gourd in southern India. Virus Dis 28:146–155 Rajeshwari R, Manasa M, Swarnalatha P, Venkataravanappa V, Krishna Reddy M (2016) Molecular detection and characterization of begomovirus associated with Ridge gourd yellow mosaic virus disease in South India. In: International Conference (VIROCON 2016), Bangalore (India). p 142 Revill PA, Ha CV, Porchun SC, Vu MT, Dale JL (2003) The complete nucleotide sequence of two distinct geminiviruses infecting cucurbits in Vietnam. Arch Virol 148(8):1523–1541 Sharma P, Verma RK, Mishra R, Sahu AK, Choudhary DK, Gaur RK (2014) First report of cucumber green mottle mosaic virus association with the leaf green mosaic disease of a vegetable crop, Luffa acutangula L. Acta Virol 58(3):299–300 Tiwari AK, Snehi SK, Singh R, Raj SK, Rao GP, Sharma PK (2011) Molecular identification and genetic diversity among six begomovirus isolates affecting cultivation of cucurbitaceous crops in Uttar Pradesh, India. Arch Phytopathol Plant Prot 45:62–72 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

L Luffa cylindrica (Sponge gourd, Suakwa) Synonyms Luffa aegyptiaca Family: Cucurbitaceae

Vegetable

Cotton leaf curl Kokhran virus

(CLCuKoV)

Synonyms Cotton leaf curl Burewala virus (CLCuBuV) Taxonomic position Genus: Begomovirus

Family: Geminiviridae

CLCuBuV infection in plants of Luffa cylindrica was reported from Pakistan (Zia-Ur-Rehman et al. 2013). The virus-infected sponge gourd plants exhibit leaf curling and stunting symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. The virus is not transmissible by mechanical inoculation nor by contact between plants. For more details of CLCuKoV, refer to Gossypium spp.

1412

Luffa cylindrica (Sponge gourd, Suakwa)

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

CGMMV infection in plants of Luffa cylindrica was reported from Pakistan and Taiwan (Hseu et al. 1987; Ali et al. 2014). The virus is not transmitted by any insect vector. The virus is transmitted through soil and irrigation water contaminated with infected plant debris. The virus is pollen transmitted and also by contact between plants. For more details of CGMMV, refer to Cucumis sativus.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Luffa cylindrica was reported from China (Huang et al. 1987; Xueping et al. 1995). The virus-infected sponge gourd plants exhibit severe mosaic, mottling, distortion, and wrinkling of leaves. Older leaves develop chlorosis and necrosis along the margins which later spread over the entire leaf (Zakaria 1999). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(CABYV)

Family: Luteoviridae

CABYV infection in plants of Luffa cylindrica was reported from China, Taiwan, and India (Xiang et al. 2008; Knierim et al. 2010; Suveditha et al. 2016). The virus-infected sponge gourd plants initially exhibit chlorotic lesions, followed by yellowing symptoms and thickening of older leaves. The virus is transmitted by aphids Myzus persicae, M. euphorbiae, and Aphis gossypii in a persistent, circulative, non-propagative manner, (Lecoq et al. 1992). The virus is not transmissible by mechanical sap-inoculation. For more details of CABYV, refer to Cucumis melo.

Papaya ringspot virus Taxonomic position Genus: Potyvirus

(PRSV)

Family: Potyviridae

PRSV pathotype W was identified in a commercial field of Luffa cylindrica in Pune, India (Verma et al. 2006). The infected sponge gourd plants showed mosaic mottling and vein-banding symptoms. A number of aphid vectors transmit this virus in a non-persistent manner; PRSV is also transmissible by mechanical sap-inoculation. For more details of PRSV, refer to Carica papaya.

Luffa cylindrica (Sponge gourd, Suakwa)

1413

Suakwa aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(SABYV)

Family: Luteoviridae

Geographical distribution SABYV infection in plants of Luffa cylindrica was reported from China, the Philippines, Thailand, Indonesia, and Taiwan (Knierim et al. 2010, 2012; Shang et al. 2009). Symptoms and host(s) The virus-infected sponge gourd plants exhibit leaf interveinal yellowing symptoms. Transmission The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not mechanically sap-transmissible. Virion properties and genome The virions are 25–30 nm in diameter, are hexagonal in outline, and have no envelope. The genome consists of one positive-sense single-stranded RNA of 5843 nt (JQ700308 = NC_018571; KX122023) (Maina et al. 2016).

L Tomato leaf curl New Delhi virus Taxonomic position Genus: Begomovirus

(ToLCNDV)

Family: Geminiviridae

The virus-infected plants of Luffa cylindrica were reported from India (Sohrab et al. 2003, 2013; Tiwari et al. 2011, 2012; Khan et al. 2014). In the early stage of the disease, the virus-infected sponge gourd plants exhibit mosaic and yellow spot on newly emerged leaves; later chlorosis and curling of the leaves will develop. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, nonpropagative manner, (Castillo et al. 2011). The virus is easily transmissible by sap to ridge gourd (Luffa acutangula), sponge gourd, and Nicotiana benthamiana (Sohrab et al. 2013). The virus is grafttransmissible. For more details of ToLCNDV, refer to Solanum lycopersicum.

References Ali A, Hussain A, Ahmad M (2014) Occurrence and molecular characterization of Cucumber green mottle mosaic virus in cucurbit crops of KPK, Pakistan. Braz J Microbiol 45:1247–1253 Castillo JN, Fiallo-Olive E, Sanchez-Campos S (2011) Emerging virus diseases transmitted by whiteflies. Annu Rev Phytopathol 49:219–248 Hseu SH, Huang CH, Chang CA, Yang WZ, Chang YM, Hsiao CH (1987) The occurrence of five viruses in six cucurbits in Taiwan. Plant Prot Bull 29:233–244 Huang CH, Chao YJ, Chang CA, Hseu SH, Hsaio CH (1987) Identification and comparison of different viruses on symptom expression in Luffa. J Agric Res China 36:413–420 Khan MS, Tiwari AK, Raj SK, Srivastava A, Hye Ji S, Chun SC (2014) Molecular epidemiology of begomoviruses occurring on some vegetables, grain legume and weed species in the Terai belt of north India. J Plant Dis Protect 121:53–57

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Lupinus spp. (Lupin)

Knierim D, Deng TC, Tsai WS, Green SK, Kenyon L (2010) Molecular identification of three distinct Polerovirus species and a recombinant Cucurbit aphid-borne yellows virus strain infecting cucurbit crops in Taiwan. Plant Pathol 59:991–1002 Knierim D, Tsai WS, Deng TC, Green S, Kenyon L (2012) Full-length genome analysis of four polerovirus isolates infecting cucurbits in Taiwan determined from total RNA extracted from field samples. NCBI. NC_018571 Lecoq H, Bourdin D, Wipf-Scheibel C, Bon M, Lot H, Lemaire O, Herrbach E (1992) A new yellowing disease of cucurbits caused by a Luteovirus, Cucurbit aphid-borne yellows virus. Plant Pathol 41:749–761 Maina S, Edwards OR, de Almeida L, Ximenes A, Jones RAC (2016) First complete genome sequence of Suakwa aphidborne yellows virus from East Timor. Genome Announc 4(4):e00718–e00716 Shang Q, Xiang H, Han C, Li D, Yu J (2009) Distribution and molecular diversity of three cucurbit-infecting poleroviruses in China. Virus Res 145:341–346 Sohrab SS, Mandal B, Pant RP, Varma A (2003) First report of association of Tomato leaf curl virus New Delhi (ToLCVNDe) with yellow mosaic disease of Luffa cylindrica in India. Plant Dis 87:1148 Sohrab SS, Karim S, Varma A, Abuzenadah AM, Chaudhary AG, Mandal B (2013) Role of sponge gourd in apical leaf curl disease of potato in Northern India. Phytoparasitica 41:403 Suveditha S, Jalali S, Krishna Reddy M (2016) Identification of Cucurbit-aphid borne yellows virus in sponge gourd (Luffa aegyptiaca) in India. In: International conference (VIROCON 2016), Bangalore (India). p 127 Tiwari AK, Snehi SK, Singh R, Raj SK, Rao GP, Sharma PK (2011) Molecular identification and genetic diversity among six Begomovirus isolates affecting cultivation of cucurbitaceous crops in Uttar Pradesh, India. Arch Phytopathol Plant Protect 45:62–72 Tiwari AK, Snehi SK, Khan MS, Sharma PK, Raj SK, Rao GP (2012) Molecular detection and identification of Tomato leaf curl New Delhi virus associated with yellow mosaic and leaf curl disease of Luffa cylindrica in India. Indian Phytopathol 65(1):80–84 Verma R, Baranwal VK, Prakash S, Tomer SPS, Pant RP, Ahlawat YS (2006) First report of Papaya ringspot virus – W in sponge gourd form India. J Phytopathol 90:974 Xiang HY, Shang QX, Han CG, Li DW, Yu JL (2008) First report on the occurrence of Cucurbit aphid-borne yellows virus on nine cucurbitaceous species in China. Plant Pathol 57:390 Xueping Z, Zhixing X, Jing X, Debao L (1995) Studies on cucumber mosaic virus isolated from Luffa cylindrica. J South Chin Agric Univ 16:74–79 Zakaria S (1999) Viruses of cucurbits. The strategies: MCB-MAPPS Plant protection conference 1999. Proceedings. pp 68–71 Zia-Ur-Rehman M, Herrmann H-W, Hameed U, Haider MS, Brown JK (2013) First detection of Cotton leaf curl Burewala virus and cognate Cotton leaf curl Multan betasatellite and Gossypium darwinii symptomless alphasatellite in symptomatic Luffa cylindrica in Pakistan. Plant Dis 97:1122

Lupinus spp. (Lupin) Family: Fabaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Grain legume

(AMV)

Family: Bromoviridae

AMV infection in plants of Lupinus spp. was reported from the UK, Germany, Poland, and Australia (Hull 1968; Jones and McLean 1989; Alberts et al. 1996). The virus-infected lupin plants exhibit tip necrosis and progressive dieback symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also through mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Lupinus spp. (Lupin)

1415

Bean common mosaic virus Taxonomic position Genus: Potyvirus

(BCMV)

Family: Potyviridae

BCMV infection in plants of Lupinus luteus was reported from Poland and Australia (Frencel and Pospieszny 1979; Jones and McLean 1989). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is seed-transmitted in Lupinus luteus seeds up to 16% (Frencel and Pospieszny 1979). For more details of BCMV, refer to Phaseolus vulgaris.

Bean yellow mosaic virus

(BYMV)

Synonyms White lupin mosaic virus

Taxonomic position Genus: Potyvirus

Family: Potyviridae

BYMV infection in plants of Lupinus spp. was reported from the UK, Poland, Hungary, the USA, Ukraine, and Australia (Hull 1968; Beczner 1970; Moskovets et al. 1971; Frencel and Pospieszny 1977; Hampton et al. 1992; Robertson and Coyne 2009; Kehoe et al. 2014a). The virus probably occurs wherever lupins are grown (Cheng and Jones 1999, 2000). The virus-infected lupin plants show yellowing and mottling of leaves followed by the formation of many small leaves near the top of the plant and curling over the stem into the form of a shepherd’s crook, necrosis and then death (Gondran et al. 1994). In some cultivars symptom severity varies, but may be little more than vein-clearing and mottle (Piche et al. 1993). Two strains of BYMV are reported, viz., the common necrotic (BYMV-N) strain kills the infected plant, and the less abundant non-necrotic (BYMV-NN) strain causes stunting without killing the plant. The virus is transmitted by more than 20 aphid species in a non-persistent manner (Berlandier et al. 1997). The virus is sap-transmissible to the members of several plant families. The virus is seed-borne in lupin seeds (Hull 1968). For more details of BYMV, refer to Phaseolus vulgaris.

Bidens mottle virus Taxonomic position Genus: Potyvirus

(BiMoV)

Family: Potyviridae

BiMoV infection in plants of Lupinus luteus was reported from Florida, USA (Edwardson et al. 1976; Jones and McLean 1989). The virus-infected lupin plants exhibit mottling and stunting symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BiMoV, refer to Bidens spp.

L

1416

Lupinus spp. (Lupin)

Clover yellow vein virus Taxonomic position Genus: Potyvirus

(ClYVV)

Family: Potyviridae

ClYVV infection in plants of Lupinus spp. was reported from Europe, Australia, and the USA (Jones and McLean 1989). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ClYVV, refer to Trifolium spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV-infected plants of Lupinus spp. were reported from the UK, Australia, Russia, Belarus, Ukraine, Lithuania, and Poland (Hull 1968; Bowyer and Keirnan 1981; Frencel and Pospieszny 1985; Samuitiene and Navalinskiene 2008). The virus-infected plants exhibit chlorosis, faint mottle, bunching and down curling of young leaves, reduction in leaf size, and brown streaking of the stem. Late infection may result in little more than chlorosis of the youngest leaves. With L. angustifolius symptoms are similar but somewhat less severe and without stem streaking (Frencel and Pospieszny 1985; Jones and McLean 1989; Wahyuni and Francki 1992). The virus is mechanically sap-transmissible and has very wide host range. It is transmitted by different aphid species: Myzus persicae, Acyrthosiphon kondoi, Aphis craccivora, and Lipaphis erysimi in a non-persistent manner (Berlandier et al. 1997). The virus is seed-transmitted in Lupinus angustifolius between 12% and 18% (Jones 1988; Jones and Proudlove 1991; Wylie et al. 1993; Geering and Randles 1994; O’Keefe et al. 2007). For more details of CMV, refer to Cucumis sativus.

Hardenbergia mosaic virus Taxonomic position Genus: Potyvirus

(HarMV)

Family: Potyviridae

HarMV infection in plants of Lupinus angustifolius and L. cosentinii was reported from Australia (Webster et al. 2007; Luo et al. 2011; Kehoe et al. 2014b). The virus-infected L. angustifolius plants exhibit symptoms of stunting, necrotic stem streaking, and tip wilting, sometimes leading to plant death, whereas L. cosentinii plants exhibit mosaic and leaf deformation symptoms. The virus is transmitted by an aphid vector, Myzus persicae, in a non-persistent manner and also by mechanical sap-inoculation. The virus is not seed-transmitted in L. angustifolius. For more details of HarMV, refer to Hardenbergia spp.

Lettuce necrotic yellows cytorhabdovirus Taxonomic position Genus: Cytorhabdovirus

Family: Rhabdoviridae

(LNYV)

Lupinus spp. (Lupin)

1417

LNYV infection in plants of Lupinus spp. was reported from Australia (Jones and McLean 1989). The virus-infected plants exhibit interveinal chlorosis and stunting symptoms. The virus is transmitted by aphid vectors in a persistent, propagative manner, and also by mechanical sap-inoculation. For more details of LNYV, refer to Lactuca sativa.

Lupinus mosaic virus Taxonomic position Genus: Potyvirus

(LuMV)

Family: Potyviridae

Geographical distribution LuMV infection in plants of Lupinus polyphyllus was reported from the south of the Czech Republic (Sarkisova and Petrzik 2009, 2011). Symptoms and host(s) The virus-infected lupin plants exhibit mild mosaic and interveinal yellowing symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically sap-transmissible to Nicotiana benthamiana, Chenopodium quinoa, and L. polyphyllus plants. Virion properties and genome The virions are non-enveloped, flexuous filaments 690 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 10,113 nt (EU847625 = NC_014898) (Sarkisova and Petrzik 2011; Revers and Garcia 2015; Wylie et al. 2017).

Nootka lupine vein clearing virus Taxonomic position Genus: Alphacarmovirus

(NLVCV)

Family: Tombusviridae

Geographical distribution NLVCV was reported for the first time on wild lupin (Lupinus nootkatensis) in south central Alaska (USA) (Robertson 2004; Robertson et al. 2007). Symptoms and host(s) The infected wild lupin plants displayed prominent leaf vein-clearing and mosaic in leaves of older plants, and in seedlings, respectively. Leaf symptoms of pronounced vein-clearing and mosaic were produced on 3- to 4-week-old plants in late June at south central Alaska (Robertson 2004). Transmission The virus is mechanically sap-transmissible, and only species belonging to the Fabaceae are susceptible to NLVCV (Robertson 2004).

L

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Lupinus spp. (Lupin)

Virion properties and genome The virions are isometric 30 nm in diameter and have T = 3 icosahedral symmetry. The genome consists of a single molecule of linear, positive-sense ssRNA of 4172 nt (EF207438 = NC_009017) (Robertson et al. 2007).

Pea early-browning virus Taxonomic position Genus: Tobravirus

(PEBV)

Family: Virgaviridae

PEBV infection in plants of Lupinus spp. was reported from Poland and Australia (Pospieszny and Frencel 1985; Jones and McLean 1989). The virus-infected lupin plants exhibit interveinal chlorosis and slight stunting symptoms. The virus is transmitted by nematode vectors, and the virus is also mechanically sap-transmissible. For more details of PEBV, refer to Pisum sativum.

Peanut mottle virus Taxonomic position Genus: Potyvirus

(PeMoV)

Family: Potyviridae

PeMoV infection in plants of Lupinus spp. was reported from the USA and Australia (Demski et al. 1981; Jones and McLean 1989). The virus-infected lupin plants exhibit mild vein-clearing, bright veinclearing, and leaf curling symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PeMoV, refer to Arachis hypogaea.

Peanut stunt virus Taxonomic position Genus: Cucumovirus

(PSV)

Family: Bromoviridae

PSV infection in plants of Lupinus spp. was reported from Poland and Australia (Jones and McLean 1989). The virus-infected lupin plants exhibit mosaic and mottling of leaves and stunting of plants. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PSV, refer to Arachis hypogaea.

Soybean dwarf virus Taxonomic position Genus: Luteovirus

(SbDV)

Family: Luteoviridae

SbDV infection in plants of Lupinus spp. was reported from Australia, Tasmania, and New Zealand (Jones and McLean 1989). The virus is transmitted by aphid vectors in a circulative, non-propagative

Lupinus spp. (Lupin)

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manner. The virus is not transmissible by mechanical inoculation. The virus is not transmissible by contact between plants. For more details of SbDV, refer to Glycine max.

Soybean mosaic virus Taxonomic position Genus: Potyvirus

(SMV)

Family: Potyviridae

SMV infection in plants of Lupinus albus was reported from Pretoria (South Africa) (Vroon et al. 1988). The virus-infected lupin plants exhibit dark green islands on leaves, leaf malformation, and internode shortening. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is seed-transmitted to the extent of 1.2% in Lupinus albus (Vroon et al. 1988). For more details of SMV, refer to Glycine max.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Lupinus spp. was reported from Australia (Sharman et al. 2015). The virus-infected lupin plants exhibit leaf deformation and tip necrosis symptoms. The virus is transmitted by the thrips vectors; the virus present in/on pollen entering the host through injuries caused by thrips while feeding. The virus is mechanically sap-transmissible but is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Tomato black ring virus Taxonomic position Genus: Nepovirus

(TBRV)

Family: Secoviridae

TBRV infection in plants of Lupinus spp. was reported from Germany, Poland, and Australia (Jones and McLean 1989). The virus is transmitted by the nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TBRV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Lupinus spp. was reported from the USA and Australia (Jones and McLean 1989; Schuster and Halliwell 1994). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

L

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Lupinus spp. (Lupin)

References Alberts EV, Nutter FW Jr, Corbett AJ, Graetz DK (1996) First report of Alfalfa mosaic virus in lupins in Australia. Plant Dis 80:1302 Beczner L (1970) Narrow-leaf virus disease of yellow Lupin. Novenyvedelmi Kutato Intezet Kozlemenyei 4:5–18 Berlandier FA, Thackray DJ, Jones RAC, Latham LJ, Cartwright L (1997) Determining the relative roles of different aphid species as vectors of cucumber mosaic and bean yellow mosaic viruses in lupins. Ann Appl Biol 131:297–314 Bowyer JW, Keirnan E (1981) Cucumber mosaic virus in lupin. Australas Plant Pathol 10:27–29 Cheng Y, Jones RAC (1999) Distribution and incidence of the necrotic and nonnecrotic strains of bean yellow mosaic virus in wild and crop lupins. Aust J Agric Res 50:589–599 Cheng Y, Jones RAC (2000) Biological properties of necrotic and non-necrotic strains of Bean yellow mosaic virus in cool season grain legumes. Ann Appl Biol 136:215–227 Demski JW, Kahn MA, Wells HD, Miller JD (1981) Peanut mottle virus in forage legumes. Plant Dis 65:359–362 Edwardson JR, Purcifull DE, Christie RG, Christie SR (1976) Blue lupine, a natural host for Bidens mottle virus. Plant Dis Reptr 60:776 Frencel I, Pospieszny H (1977) Viruses in natural infection of yellow lupin (Lupinus luteus L.) in Poland. I. Bean yellow mosaic virus. Acta Phytopathol Acad Sci Hung 12:87–90 Frencel I, Pospieszny H (1979) Viruses in natural infections of yellow lupin (Lupinus luteus L.) in Poland IV. Bean common mosaic virus (BCMV). Acta Phytopathol Acad Sci Hung 14:279–284 Frencel I, Pospieszny K (1985) Viruses and natural infections of yellow lupin (Lupinus luteus L.) in Poland. V. Cucumber mosaic virus. Acta Phytopathol Acad Sci Hung 20:87–90 Geering ADW, Randles JW (1994) Interactions between a seed-borne strain of cucumber mosaic cucumovirus and its lupin host. Ann Appl Biol 124:301–314 Gondran J, Bournovalle R, Duthion C (1994) Identification of diseases, pests and physical constraints in white lupin. INRA, Versailles 48 pp Hampton RO, Shukla DD, Jordan RL (1992) Comparative potyvirus host range, serology and coat protein peptide profiles of White lupin mosaic virus. Phytopathology 82:566–571 Hull R (1968) Virus diseases of garden lupin in Great Britain. Ann Appl Biol 61:373–380 Jones RAC (1988) Seed borne Cucumber mosaic virus infection of narrow-leafed lupin (Lupinus angustifolius) in Western Australia. Ann Appl Biol 113:507–518 Jones RAC, McLean GD (1989) Virus diseases of lupins. Ann Appl Biol 114:609–637 Jones RAC, Proudlove W (1991) Further studies on Cucumber mosaic virus infection of narrow-leafed lupin (Lupinus angustifolius): seed-borne infection, aphid transmission, spread and effects on grain yield. Ann Appl Biol 118:319–329 Kehoe MA, Buirchell BJ, Coutts BA, Jones RAC (2014a) Black pod syndrome of Lupinus angustifolius is caused by late infection with Bean yellow mosaic virus. Plant Dis 98:739–745 Kehoe MA, Coutts BA, Buirchell BJ, Jones RAC (2014b) Hardenbergia mosaic virus: crossing the barrier between native and introduced plant species. Virus Res 184:87–92 Luo H, Wylie SJ, Coutts B, Jones RAC, Jones MGK (2011) A virus of an isolated indigenous flora spreads naturally to an introduced crop species. Ann Appl Biol 159:339–347 Moskovets SM, Krayev VG, Porembs’ka NB, Bilik LG, Baratova DF (1971) Kiev ‘Naukova Dumka’, viruses and virus diseases of legumes in the Ukraine. 136 pp O’Keefe DC, Berryman DI, Coutts BA, Jones RAC (2007) Lack of seed coat contamination with Cucumber mosaic virus in lupin permits reliable, large-scale detection of seed transmission in seed samples. Plant Dis 91:504–508 Piche C, Peterson J, Fortin MG (1993) A note on the detection of Bean yellow mosaic virus infecting white lupine in Canada. Phytoprotection 74:153–155 Pospieszny H, Frencel I (1985) Viruses in natural infection of yellow lupin (Lupinus luteus L.) in Poland. VI. Pea early browning virus. Acta Phytopathol Acad Sci Hung 20:91–95 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Robertson NL (2004) Biology of a new virus isolated from Lupinus nootkatensis plants in Alaska. Plant Pathol 53(5):569–576 Robertson NL, Coyne CJ (2009) First report of Bean yellow mosaic virus from diseased Lupinus luteus in Eastern Washington. Plant Dis 93:319 Robertson NL, Cote F, Pare C, Leblanc E, Bergeron MG, Leclerc D (2007) Complete nucleotide sequence of Nootka lupine vein-clearing virus. Virus Genes 35:807–814 Samuitiene M, Navalinskiene M (2008) Occurrence of Cucumber mosaic cucumovirus on ornamental plants in Lithuania. Zemdirbyste-Agriculture 95:135–143 Sarkisova T, Petrzik K (2009) A new potyvirus identified in Czech Republic. Acta Virol 53:143 Sarkisova T, Petrzik K (2011) Determination of the complete nucleotide sequence of a lupine potyvirus isolate from the Czech Republic reveals that it belongs to a new member of the genus Potyvirus. Arch Virol 156:167–169

Lycianthes rantonnetii (Blue potato bush)

1421

Schuster GL, Halliwell RS (1994) Six new hosts of tomato spotted wilt virus in Texas. Plant Dis 78:100 Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct Tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207 Vroon CW, Pietersen C, van Tonder HJ (1988) Seed transmission of Soybean mosaic virus in Lupinus albus L. Phytophylactica 20:169–175 Wahyuni WS, Francki RIB (1992) Responses of some grain and pasture legumes to 16 strains of Cucumber mosaic virus (CMV). Aust J Agric Res 43:465–477 Webster CG, Coutts BA, Jones RAC, Jones MGK, Wylie SJ (2007) Virus impact at the interface of an ancient ecosystem and a recent agroecosystem: studies on three legume-infecting potyviruses in the southwest Australian floristic region. Plant Pathol 56:729–742 Wylie S, Wilson CR, Jones RAC, Jones MGK (1993) A polymerase chain reaction assay for Cucumber mosaic virus in lupin seeds. Aust J Agric Res 44:41–51 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Lycaste spp. Family: Orchidaceae

Ornamental

Cymbidium mosaic virus Taxonomic position Genus: Potexvirus

(CymMV)

L Family: Alphaflexiviridae

CymMV infection in plants of Lycaste spp. was reported from Costa Rica (Arce-Rodriguez et al. 2014). The virus-infected lycaste plants exhibit symptoms of yellow-green mosaic pattern on both surfaces of leaves. Often the entire leaf shows this pattern but occasionally only part of the leaf is infected. The virus is mechanically sap-transmissible. The virus is also transmissible by contact between infected and healthy plants, and also spreads through contact by contaminated cutting tools, equipment, human hands, potting media, or water. No vector transmission is reported. For more details of CymMV, refer to Cymbidium spp.

References Arce-Rodriguez A, Montero-Carmona W, Varela I, Gätjens-Boniche O (2014) Detection of Cymbidium mosaic virus (CymMV) and Odontoglossum ringspot virus (ORSV) in orchids grown in Costa Rica. Biotecnia XVI(3):3–10

Lycianthes rantonnetii (Blue potato bush) Synonyms Lycianthes biflora; Solanum rantonnetii Family: Solanaceae

Ornamental

1422

Lycianthes rantonnetii (Blue potato bush)

Citrus exocortis viroid Taxonomic position Genus: Pospiviroid

(CEVd)

Family: Pospiviroidae

CEVd infection in plants of Lycianthes rantonnetii was reported from Italy (Luigi et al. 2011). The viroid-infected blue potato bush plants do not exhibit any symptoms. The viroid is mechanically saptransmissible and also transmissible through contaminated tools. Use of infected budwood and root stocks is the primary mode of spread. For more details of CEVd, refer to Citrus spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Lycianthes rantonnetii was reported from Hungary (Salamon et al. 2013). The virus-infected blue potato bush plants exhibit yellow-green mosaic symptoms on the top leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Lycianthes yellow mosaic virus Taxonomic position Genus: Begomovirus

(LyYMV)

Family: Geminiviridae

Geographical distribution LyYMV infection in plants of Lycianthes biflora was reported from China (Tang et al. 2017). Symptoms and host(s) The virus-infected lycianthes plants exhibit yellow mosaic symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2752 nt (KT582302 = NC_038978) and DNA-B of 2704 nt (KT582303) (Briddon 2001; Brown et al. 2015; Tang et al. 2017; Zerbini et al. 2017).

Potato spindle tuber viroid Taxonomic position Genus: Pospiviroid

(PSTVd)

Family: Pospiviroidae

Lycianthes rantonnetii (Blue potato bush)

1423

PSTVd infection in plants of Lycianthes rantonnetii was reported from Croatia and Italy (Di Serio 2007; Luigi et al. 2011; Matousek et al. 2014; Milanovic et al. 2014). The viroid is mechanically sap-transmissible. Use of infected budwood and root stocks is the primary mode of spread. For more details of PSTVd, refer to Solanum tuberosum.

Tomato apical stunt viroid Taxonomic position Genus: Pospiviroid

(TASVd)

Family: Pospiviroidae

TASVd infection in plants of Lycianthes rantonnetii was reported from the Netherlands and Belgium (Verhoeven et al. 2010, 2012; Hennig et al. 2013; Milanovic et al. 2014; Van Bogaert et al. 2017). The viroid-infected plants do not exhibit any symptoms. The L. rantonnetii isolate is mechanically sap-transmissible. For more details of TASVd, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Solanum rantonnetii was reported from Italy (Dellavalle et al. 1999). The virus-infected blue potato bush plants exhibit necrotic spot and ring-spot symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Briddon RW (2001) Begomovirus. Geminiviridae. The Springer index of plant viruses. Springer, New York, pp 340–348, https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Dellavalle G, Ciuffo M, Roggero P, Lisa V, Minuto A, Minuto G, Rapetti S (1999) Tospoviruses in Delphinium sp., Gazania, Marguerite, Celery, Tragopogon porrifolius and Solanum rantonnetti in Liguria (Northern Italy). Inf Fitopatol 49:63–64 Di Serio F (2007) Identification and characterization of potato spindle tuber viroid infecting Solanum jasminoides and S. rantonnetii in Italy. J Plant Pathol 89:297–300 Hennig E, Pięcinska J, Borodynko N, Hasiow-Jaroszewska B (2013) First reports of Potato spindle tuber viroid (PSTVd) on Solanum jasminoides and of Tomato apical stunt viroid (TASVd) on Solanum rantonnetti in Poland. Plant Dis 97:1663 Luigi M, Luison D, Tomassoli L, Faggioli F (2011) Natural spread and molecular analysis of pospiviroids infecting ornamentals in Italy. J Plant Pathol 93:491–495 Matousek J, Piernikarczyk RJJ, Dedic P, Mertelík J, Uhlířová K, Duraisamy G, Orctová L, Kloudová K, Ptáček J, Steger G (2014) Characterization of Potato spindle tuber viroid (PSTVd) incidence and new variants from ornamentals. Eur J Plant Pathol 138:93–101 Milanovic J, Kajic V, Mihaljevic S (2014) Occurrence and molecular variability of Potato spindle tuber viroid and Tomato apical stunt viroid in ornamental plants in Croatia. Eur J Plant Pathol 139:785–788

L

1424

Lycium barbarum (Chinese wolfberry)

Salamon P, Nemes K, Salánki K, Palkovics L (2013) First record of virus infection of blue potato bush, Lycianthes rantonnetii. J Plant Pathol 95:S4.74 Tang YF, He ZF, Brown JK, She XM, Lan GB (2017) Molecular characterization of a novel bipartite Begomovirus isolated from Lycianthes biflora in China. Arch Virol 162(8):2473–2476 Van Bogaert N, Smagghe G, Maes M, De Backer M, De Jonghe K (2017) Phylogeny of five predominant pospiviroid species in Belgium. Eur J Plant Pathol 149:25–33 Verhoeven JTJ, Botermans M, Jansen CCC, Roenhorst JW (2010) First report of Tomato apical stunt viroid in the symptomless hosts Lycianthes rantonnetii and Streptosolen jamesonii in the Netherlands. Plant Dis 94(6):791 Verhoeven JTJ, Botermans M, Meekes ETM, Roenhorst JW (2012) Tomato apical stunt viroid in the Netherlands: most prevalent pospiviroid in ornamentals and first outbreak in tomatoes. Eur J Plant Pathol 133(4):803–810 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Lycium barbarum (Chinese wolfberry) Family: Solanaceae

Medicinal

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Lycium barbarum was reported from East Germany and Bulgaria by Schmelzer and Schmidt (1968). The virus-infected Chinese wolfberry plants exhibit symptoms of chlorotic mottling with some necrosis. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

References Schmelzer K, Schmidt HE (1968) Untersuchungen an Viren der Zier-und Wildegeholze. 6. Mitteilung: Erganzende Befunde an Caryopteris sowie Virosen an Philadelphus, Aristolochia, Buddleja, Lycium und Aesculus. Phytopathol Z 62:105–126

Lycoris spp. (Spider lily) Family: Amaryllidaceae

Ornamental

Lycoris mild mottle virus Taxonomic position Genus: Potyvirus

(LyMMoV)

Family: Potyviridae

Lycoris spp. (Spider lily)

1425

Geographical distribution LyMMoV infection in plants of golden spider lily (Lycoris aurea) was reported from Taiwan (Chang et al. 2002). Symptoms and host(s) The virus-infected spider lily plants exhibit mild mottle symptoms. Transmission The virus is transmitted by aphid vectors in a non-persistent manner. The virus is also mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, flexuous filaments, 750 nm long and 11–13 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA of 9.5–10 kb (Revers and Garcia 2015). A partial genome sequence of 1989 nt is available (AF399672) (Revers and Garcia 2015; Wylie et al. 2017).

Narcissus degeneration virus

(NDV)

L

Synonyms Lycoris potyvirus Taxonomic position Genus: Potyvirus

Family: Potyviridae

NDV infection in plants of golden spider lily (Lycoris aurea) was reported from Taiwan (Chang et al. 2002). The virus-infected spider lily plants exhibit severe mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of NDV, refer to Narcissus spp.

Nerine latent virus

(NeLV)

Synonyms Lycoris virus T Taxonomic position Genus: Carlavirus

Family: Betaflexiviridae

NeLV infection in plants of Lycoris aurea was reported from Taiwan (Chen et al. 2005). The virusinfected spider lily plants exhibit foliar streaking or mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of NeLV, refer to Nerine spp.

1426

Lysimachia spp.

References Chang C-A, Chen C-C, Hsu H-T (2002) Partial characterization of two potyviruses associated with Golden spider lily severe mosaic disease. Acta Hortic 568:127–134 Chen C-C, Chang F-L, Cheng Y-H, Chang Y-M, Lin Y-Y, Chang C-A (2005) Lycoris virus T, a carlavirus newly identified in golden spider lily also found widespread in narcissus and amaryllis. Plant Pathol Bull 14:300 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Lysimachia spp. Family: Primulaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Lysimachia spp. was reported from Spain (Albouy and Devergne 1999). The virus-infected lysimachia plants exhibit mosaic and leaf distortion symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in Lysimachia congestiflora was reported from the Netherlands (Werkman et al. 2007). The virus-infected lysimachia plants exhibit symptoms of chlorotic mottle and may also show necrotic leaf margins. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Lysimachia spp. was reported from the Netherlands (Werkman et al. 2007). The virus-infected lysimachia plants exhibit symptoms of chlorotic mottle and may also show necrotic leaf margins. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Lysimachia spp.

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References Albouy J, Devergne JC (1999) Diseases produced by viruses on ornamental plants. Ediciones Mundi-Prensa, Madrid. 480 p Werkman AW, Verhoeven JThJ, Roenhorst JW (2007) Plant species found infected by Tomato spotted wilt virus and Impatiens necrotic spot virus at the Dutch Plant Protection Service since 1989. www.tuinbouw.nl/sites/default/files/ 14761.19%20Bijlage%201%20NL%20viruslijst%20TSWV-INSV%202007.pdf

L

M

Maclura pomifera (Osage orange) Family: Moraceae

Maclura mosaic virus Taxonomic position Genus: Macluravirus

Trees/Shrubs

(MacMV)

Family: Potyviridae

Geographical distribution MacMV infection in plants of Maclura pomifera spreads in the former Yugoslavia (Plese and Milicic 1973; Plese and Stefanac 1976; Plese et al. 1979). Symptoms and host(s) With MacMV infection of M. pomifera, the symptoms consist of yellow mosaic associated with foliar distortion and interveinal yellowing which in turn causes local tissue translucency, which is more obvious in spring than during summer periods of high temperature (Plese and Milicic 1973; Plese and Stefanac 1976). Transmission The virus is transmitted by an aphid vector Myzus persicae in a non-persistent manner. The virus is also mechanically sap-transmissible to several families; local lesions are produced on Chenopodium species. The virus is also transmissible through grafting. Virion properties and genome The virions are non-enveloped, flexuous filaments, with a clear modal length of 672 nm and 13–16 nm wide. The genome consists of a single molecule of linear positive-sense ssRNA of c.9.5 kb. A partial genome sequence of 2413 nt is available (U58771) (Plese et al. 1979; Badge et al. 1997; Lopez-Moya et al. 2009; Foster 2011; Wylie et al. 2017). © Springer Nature India Private Limited 2019 K. S. Sastry et al., Encyclopedia of Plant Viruses and Viroids, https://doi.org/10.1007/978-81-322-3912-3

1430

Macroptilium spp.

References Badge J, Robinson DJ, Brunt AA, Foster GD (1997) 30 -Terminal sequences of the RNA genomes of narcissus latent and Maclura mosaic viruses suggest that they represent a new genus of the Potyviridae. J Gen Virol 78:253–257 Foster GD (2011) Macluravirus. Potyviridae. In: The Springer Index of Viruses. Springer, New York, pp 1421–1424. https://doi.org/10.1007/978-0-387-95919-1_234 Lopez-Moya JJ, Valli A, Garcia JA (2009) Potyviridae. In: Encyclopaedia of Life Sciences (ELS). Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000755.pub2 Plese N, Milicic D (1973) Two viruses isolated from Maclura pomifera. Phytopathol Z 77:178–183 Plese N, Stefanac Z (1976) Some properties of Maclura mosaic virus a member of the potyvirus group. Mitt Biol Bund Anst Forstw 170:47–50 Plese M, Koenig R, Lesemann D-E, Bozarth RF (1979) Maclura mosaic virus – an elongated plant virus of uncertain classification. Phytopathology 69:471–475 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV Virus Taxonomy Profile: Potyviridae. J Gen Virol 98:352–354

Macroptilium spp. Family: Fabaceae

Forage crop

Bean common mosaic virus Taxonomic position Genus: Potyvirus

(BCMV)

Family: Potyviridae

BCMV infection in plants of Macroptilium atropurpureum was reported from Western Australia (Saqib et al. 2010). The virus-infected macroptilium plants exhibit symptoms of severe leaf mosaic, leaf deformation, plant stunting, and malformation of pods. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. The virus is seed-transmitted in Macroptilium lathyroides plants from 5% to 33% (Provvidenti and Braverman 1976). For more details of BCMV, refer to Phaseolus vulgaris.

Bean golden yellow mosaic virus Taxonomic position Genus: Begomovirus

(BGYMV)

Family: Geminiviridae

BGYMV infection in plants of Macroptilium lathyroides was reported from Puerto Rico (Bracero et al. 2003). The virus-infected macroptilium plants exhibit golden yellow mosaic symptoms. The virus is naturally transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. For more details of BGYMV, refer to Phaseolus vulgaris.

Macroptilium bright mosaic virus Taxonomic position Genus: Begomovirus

(MacBMV)

Family: Geminiviridae

Macroptilium spp.

1431

Geographical distribution MacBMV infection in plants of Macroptilium lathyroides was reported from Brazil (Passos et al. unpublished; KX691400). Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2636 nt (KX691400 = NC_031452) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Macroptilium common mosaic virus Taxonomic position Genus: Begomovirus

(MacCMV)

Family: Geminiviridae

Geographical distribution MacCMV infection in plants of Macroptilium lathyroides was reported from Brazil (Passos et al. unpublished; KX691396). Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2632 nt (KX691396 = NC_031448) and DNA-B of 2598 nt (KX691412 = NC_031453) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Macroptilium golden mosaic virus Taxonomic position Genus: Begomovirus

(MacGMV)

Family: Geminiviridae

Geographical distribution MacGMV infection in plants of Macroptilium lathyroides was reported from Ceara (Brazil) and Jamaica (Lima et al. 2012). Symptoms and host(s) The virus-infected macroptilium plants exhibit golden mosaic symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. The virus is transmissible by grafting.

M

1432

Macroptilium spp.

Virion properties and genome The virions are twinned (geminate), incomplete icosahedra, and T = 1, 22  38 nm with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2605 nt (EU158096 = NC_010952), and DNA-B of 2553 nt (EU158097 = NC_010953) (Briddon 2001; Lima et al. 2012; Brown et al. 2015; Zerbini et al. 2017).

Macroptilium mosaic Puerto Rico virus Taxonomic position Genus: Begomovirus

(MacMPRV)

Family: Geminiviridae

Geographical distribution MacMPRV infection in plants of Macroptilium lathyroides was reported from the Caribbean region, Central America, and Puerto Rico (Idris et al. 1999, 2003). Symptoms and host(s) The virus-infected macroptilium plants exhibit bright yellow mosaic symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, type B in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate), incomplete icosahedra, and T = 1, 22  38 nm with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2615 nt (AY044133 = NC_004097) and DNA-B of 2571 nt (AY044134 = NC_004098) (Briddon 2001; Idris et al. 2003; Brown et al. 2015; Zerbini et al. 2017).

Macroptilium yellow mosaic Florida virus Taxonomic position Genus: Begomovirus

(MacYMFV)

Family: Geminiviridae

Geographical distribution MacYMFV infection in plants of Macroptilium lathyroides was reported from the Caribbean region, and Central America (Idris et al. 2003). Symptoms and host(s) The virus-infected macroptilium plants exhibit golden mosaic symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner.

Macroptilium spp.

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Virion properties and genome The virions are twinned (geminate), incomplete icosahedra, and T = 1, 22  38 nm with a single coat protein. The genome is a bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2642 nt (AY044135 = NC_004099) and DNA-B of 2605 nt (AY044136 = NC_004100) (Briddon 2001; Idris et al. 2003; Brown et al. 2015; Zerbini et al. 2017).

Macroptilium yellow mosaic virus Taxonomic position Genus: Begomovirus

(MacYMV)

Family: Geminiviridae

Geographical distribution MacYMV infection in plants of Macroptilium lathyroides was reported from India, Pakistan, Cuba, and Jamaica (Ramos et al. 2002; Amarakoon et al. 2008). Symptoms and host(s) The virus-infected macroptilium plants exhibit bright yellow mosaic, chlorosis, and leaf curling symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate), incomplete icosahedra, and T = 1, 22  38 nm with a single coat protein. The genome is a bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2630 nt (EF585290 = NC_010647; AJ344452) and DNA-B of 2593 nt (NC_010648) (Briddon 2001; Amarakoon et al. 2008; Brown et al. 2015; Zerbini et al. 2017).

Macroptilium yellow spot virus Taxonomic position Genus: Begomovirus

(MacYSV)

Family: Geminiviridae

Geographical distribution MacYSV infection in plants of Macroptilium lathyroides was reported from Brazil (Silva et al. 2012). Symptoms and host(s) The virus-infected macroptilium plants exhibit yellow spot symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner.

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Macroptilium spp.

Virion properties and genome The virions are twinned (geminate), incomplete icosahedra, and T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular, single-stranded DNA (DNA-A) of 2660 nt (JN419013 = NC_016999) (Briddon 2001; Silva et al. 2012; Brown et al. 2015; Zerbini et al. 2017).

Macroptilium yellow vein virus Taxonomic position Genus: Begomovirus

(MacYVV)

Family: Geminiviridae

Geographical distribution MacYVV infection in plants of Macroptilium spp. was reported from Brazil (Silva et al. 2012). Symptoms and host(s) The virus-infected macroptilium plants exhibit yellow vein symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate), incomplete icosahedra, and T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular, single-stranded DNA (DNA-A) of 2656 nt (JN419021 = NC_017000) (Briddon 2001; Silva et al. 2012; Brown et al. 2015; Zerbini et al. 2017).

Passiflora foetida virus Y

(PfVY)

Taxonomic position PfVY is a tentative member of the genus Potyvirus and family Potyviridae PfVY infection in plants of Macroptilium atropurpureum was reported from Taiwan (Chiang et al. 2012). The virus-infected macroptilium plants exhibit mosaic and leaf deformation symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of PfVY, refer to Passiflora foetida.

Sida golden mosaic virus Taxonomic position Genus: Begomovirus

(SiGMV)

Family: Geminiviridae

SiGMV infection in plants of Macroptilium lathyroides was reported from Jamaica (Roye et al. 1997). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. For more details of SiGMV, refer to Sida spp.

Macroptilium spp.

Siratro 1 virus Y

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(SVY1)

Taxonomic position SVY1 is a tentative member of the genus Potyvirus and family Potyviridae Geographical distribution SVY1 infection in plants of Macroptilium atropurpureum was reported from Australia (Wei et al. unpublished - DQ098900; Gibbs et al. 2008). Transmission Presumably the virus is transmitted by aphid vectors. Virion properties and genome The virions are flexuous filaments. The genome is a single molecule of positive-sense, single-stranded RNA. A partial genome sequence of 1720 nt is available (DQ098900) (Gibbs et al. 2008; Wylie et al. 2017).

Siratro 2 virus Y

(SVY2)

Taxonomic position SVY2 is a tentative member of the genus Potyvirus and family Potyviridae Geographical distribution SVY1 infection in plants of Macroptilium atropurpureum was reported from Australia (Wei et al. unpublished - DQ098901; Gibbs et al. 2008). Transmission Presumably the virus is transmitted by aphid vectors. Virion properties and genome The virions are flexuous filaments. The genome is a single molecule of positive-sense, single-stranded RNA. A partial genome sequence of 1705 nt is available (DQ098901) (Gibbs et al. 2008; Wylie et al. 2017).

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Macroptilium lathyroides was reported from Australia (Sharman et al. 2015). The virus-infected macroptilium plants exhibit symptoms of chlorotic mottle, line patterns, and local lesions. The virus is transmitted by the thrips vectors; the virus is present in/on pollen, entering the host through injuries caused by thrips while feeding. The virus is mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

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Macroptilium spp.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Macroptilium lathyroides was reported from South Florida (Ying and Davis 2000). The virus-infected Macroptilium plants exhibit symptoms of curling, distortion and chlorosis of leaves, stunting of the plant, and a high rate of flower abscission. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. The virus is transmissible by grafting but not by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

References Amarakoon II, Roye ME, Briddon RW, Bedford ID, Stanley J (2008) Molecular and biological characterization of Macroptilium yellow mosaic virus from Jamaica. Plant Pathol 57:417–426 Bracero V, Rivera LI, Beaver JS (2003) DNA analysis confirms Macroptilium lathyroides as alternative hosts of bean golden yellow mosaic virus. Plant Dis 87:1022–1025 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer Index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Chiang C-H, Fan Y-T, Yu T-A, Cheng Y-H, Chen YK (2012) First report of Passiflora virus Y infecting Macroptilium atropurpureum in Taiwan. Plant Dis 96:918 Gibbs AJ, Mackenzie AM, Wei KJ, Gibbs MJ (2008) The potyviruses of Australia. Arch Virol 153:1411–1420 Idris AM, Bird J, Brown JK (1999) First report of a bean-infecting begomovirus from Macroptilium lathyroides in Puerto Rico that is distinct from bean golden mosaic virus. Plant Dis 83:1071 Idris AM, Hiebert E, Bird J, Brown JK (2003) Two newly described begomoviruses of Macroptilium lathyroides and common bean. Phytopathology 93:774–783 Lima JAA, do Nascimento AKQ, Amorim VBO, Radaelli P, Filho JT (2012) Serological detection and molecular characterization of a begomovirus isolate obtained from Macroptilium lathyroides. Rev Caatinga 25:14–20 Provvidenti R, Braverman SW (1976) Seed transmission of bean common mosaic virus in phasemy bean. Phytopathology 66:1274–1275 Ramos PL, Fernandez A, Castrillo G, Diaz L, Echemendia AL, Fuentes A, Peral R, Pujol M, Ascencio-Ibanez JT, RiveraBustamante R, Arguello-Astorga G (2002) Macroptilium yellow mosaic virus, a new begomovirus infecting Macroptilium lathyroides in Cuba. Plant Dis 86:1049 Roye ME, McLaughlin WA, Nakhla MK, Maxwell DP (1997) Genetic diversity among geminiviruses associated with the weed species Sida spp., Macroptilium lathyroides, and Wissadula amplissima from Jamaica. Plant Dis 81:1251–1258 Saqib M, Nouri S, Cayford B, Jones RAC, Jones MGK (2010) Genome sequences and phylogenetic placement of two isolates of bean common mosaic virus from Macroptilium atropurpureum in North-West Australia. Australas Plant Pathol 39:184–191 Sharman M, Thomas JE, Persley DM (2015) Natural host range, thrips and seed transmission of distinct Tobacco streak virus strains in Queensland, Australia. Ann Appl Biol 167:197–207 Silva SJC, Castillo-Urquiza GP, Hora BT Jr, Assuncao IP, Lima GSA, Pio-Ribeiro G, Mizubuti ESG, Zerbini FM (2012) Species diversity, phylogeny and genetic variability of begomovirus populations infecting leguminous weeds in northeastern Brazil. Plant Pathol 61:457–567 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Ying ZT, Davis MJ (2000) Partial characterization and host range of tomato yellow leaf curl virus in South Florida. Proc Fla State Hortic Soc 113:185–190 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Macrotyloma axillare (Perennial horse gram)

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Macrotyloma axillare (Perennial horse gram) Family: Fabaceae

Fodder

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

(AMV)

Family: Bromoviridae

AMV infection in plants of Macrotyloma axillare was reported from Ethiopia (Edeme and Hanson 2000). The virus-infected perennial horse gram plants exhibit chlorotic mottling, mosaic, malformation, and leaf-rolling symptoms. The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Bean common mosaic virus Taxonomic position Genus: Potyvirus

(BCMV)

Family: Potyviridae

BCMV infection in plants of Macrotyloma axillare was reported from Ethiopia (Edeme and Hanson 2000). The virus-infected perennial horse gram plants exhibit chlorotic mottling, mosaic, malformation, and leaf-rolling symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BCMV, refer to Phaseolus vulgaris.

Broad bean stain virus Taxonomic position Genus: Comovirus

(BBSV)

Family: Secoviridae

BBSV infection in plants of Macrotyloma axillare was reported from Ethiopia (Edeme and Hanson 2000). The virus-infected perennial horse gram plants exhibit chlorotic mottling, mosaic, malformation, and leaf-rolling symptoms. The virus is transmitted by weevil vectors. The virus is mechanically sap-transmissible and also by grafting. For more details of BBSV, refer to Vicia faba.

References Edeme J, Hanson J (2000) First report of alfalfa mosaic, bean common mosaic, and broad bean stain viruses in Macrotyloma axillare. Plant Dis 84(5):594

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Macrotyloma uniflorum (Horse gram)

Macrotyloma uniflorum (Horse gram) Family: Fabaceae

Grain legume

Horsegram yellow mosaic virus Taxonomic position Genus: Begomovirus

(HgYMV)

Family: Geminiviridae

Geographical distribution HgYMV infection in plants of Macrotyloma uniflorum was first reported from India (Williams et al. 1968). The virus spreads in India and Pakistan (Muniyappa and Reddy 1976; Muniyappa et al. 1978, 1987; Nagaraj 1982; Qazi et al. 2007; Muniyappa et al. 2008; Barnabas et al. 2010; Prema 2013). Symptoms and host(s) The virus-infected horse gram plants exhibit symptoms of yellow mosaic on leaves, reduction of leaf size, and plant height in severely infected plants. The natural host range of this virus includes French bean, groundnut, lima bean, mung bean, pigeon pea, soybean, Bambara groundnut, and Indigofera hirsuta (Barnabas et al. 2010; Prema 2013). Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. The virus is not transmissible by mechanical inoculation, not transmissible by contact between plants, not transmitted by seed, and not transmitted by pollen. Virion properties and genome The virions are twinned (geminate), incomplete icosahedra, and T = 1, 22  38 nm with a single coat protein. The genome is a bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2728 nt (AJ627904 = NC_005635) and DNA-B of 2677 nt (AJ627905 = NC_005636) (Harrison et al. 1977; Briddon 2001; Qazi et al. 2007; Barnabas et al. 2010; Brown et al. 2015; Zerbini et al. 2017).

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Macrotyloma uniflorum was reported from Andhra Pradesh (India) (Seshadri Goud et al. 2013). The virus-infected horse gram plants exhibit symptoms of leaves with necrotic spots and wrinkled margins, together with plant stunting and wilting. The virus is transmitted by thrips vectors, which assist in the transfer of virus-infected pollen to the healthy plants. The virus is mechanically sap-transmissible and is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

Malachra alceifolia (Yellow leafbract)

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References Barnabas AD, Radhakrishnan GK, Ramakrishnan U (2010) Characterization of a begomovirus causing horsegram yellow mosaic disease in India. Eur J Plant Pathol 127:41–51 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer Index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Harrison BD, Barker H, Bock KR, Guthrie EJ, Meredith G, Atkinson M (1977) Plant viruses with circular single-stranded DNA. Nature 270:760–762 Muniyappa V, Reddy HR (1976) Studies on the yellow mosaic disease of horsegram (Dolichos biflorus L.)- virus vector relationships. Mysore J Agric Sci 10:605–610 Muniyappa V, Reddy HR, Mustak Ali TM (1978) Studies on the yellow mosaic disease of horsegram Dolichos biflorus, IV. Epidemiology of the disease. Mysore J Agric Sci 12:277–279 Muniyappa V, Rajeshweri R, Bharathan N, Reddy DVR, Nolt BL (1987) Isolation and characterization of a geminivirus causing yellow mosaic disease of Horsegram (Macrotyloma uniflorum (Lam.) Verdc.) in India. Indian J Phytopathol 119:81–87 Muniyappa V, Rajeshwari R, Bharathan N, Reddy DNR, Nolt BL (2008) Isolation and characterization of a geminivirus causing yellow mosaic disease of horsegram (Macrotyloma uniflorum [Lam.] Verdc.) in India. J Phytopathol 119:81–87 Nagaraj NC (1982) Studies on horsegram (Macrotyloma uniflorum) yellow mosaic disease. MSc (Agri.) thesis, University of Agricultural Sciences, Bangalore, p 125 Prema GU (2013) Molecular characterization of Horsegram yellow mosaic virus and its management. PhD thesis submitted to the Department of Plant Pathology, University of Agricultural Sciences, GKVK, Bangalore, p 271 Qazi J, Ilyas M, Mansoor S, Briddon RW (2007) Legume yellow mosaic viruses: genetically isolated begomoviruses. Mol Plant Pathol 8(4):343–348 Seshadri Goud TE, Vemana K, Reddy DL, Mahammed Khureshee CS, Padma JG, Shabbir S, Venkateswarlu NC, Naik KSS, Sampath Kumar D, Anthony Johnson AM, Subramanyam K (2013) First report of Tobacco streak ilarvirus infecting jasmine and horse gram. New Dis Rep 28:7 Williams FJ, Grewal JS, Amin KS (1968) Serious and new disease of pulse crops in India in 1966. Plant Dis Reptr 52:300–304 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Malachra alceifolia (Yellow leafbract) Family: Malvaceae

Weed host

Tobacco leaf curl Cuba virus Taxonomic position Genus: Begomovirus

(TbLCCuV)

Family: Geminiviridae

TbLCCuV infection in plants of Malachra alceifolia was reported from Jamaica (Hall et al. 2008). The virus-infected yellow leafbract plants show yellow mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. The virus is not transmissible by mechanical inoculation. For more details of TbLCCuV, refer to Nicotiana tabacum.

References Hall GC, Graham AP, Roye ME (2008) Tobacco leaf curl Cuba virus infects the weed Malachra alceifolia in Jamaica. Plant Pathol 57:398

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Malachra capitata (Brazil jute)

Malachra capitata (Brazil jute) Family: Malvaceae

Fiber crop

Cotton leaf curl Barasat virus

(CLCuBarV)

Synonyms Malachra yellow mosaic virus (MalYMV) Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution MalYMV infection in plants of Malachra capitata was reported from Barasat, West Bengal (India) (Roy and Sherpa 2018). Symptoms and host(s) The virus-infected Malachra capitata plants show leaf curling, yellow mosaic, vein clearing, leaf yellowing and stunted growth of the plants. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2739 nt (LC080677 = NC_039003) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017; Roy and Sherpa 2018).

References Briddon RW (2001) Begomovirus. Geminiviridae. The Springer index of plant viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Roy B, Sherpa AR (2018) Detection and molecular characterization of a new Begomovirus associated with mosaic disease of Malachra capitata (Malvaceae). Aust Plant Dis Notes 13:30 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Malus domestica (Apple) Family: Rosaceae

Edible fruit

Apple chlorotic leaf spot virus Taxonomic position Genus: Trichovirus

(ACLSV)

Family: Betaflexiviridae

Malus domestica (Apple)

1441

Geographical distribution ACLSV was first reported in plants of Malus spp. from the USA (Mink and Shay 1959, 1962; Lister et al. 1965). This virus is distributed worldwide, probably wherever apples are cultivated: Eastern Asian region, the Eurasian region, the North American region, the Pacific region, India, Iran, Australia, China, Japan, Ethiopia, the Czech Republic, Korea, Turkey, and New Zealand (Cropley et al. 1963; Cropley 1968; Chairez and Lister 1973; Fridlund 1973; Flegg and Clark 1979; Barba and Clark 1986; Delbos and Dunez 1988; Li and Mink 1989; Cieslinska et al. 1995; Kinard et al. 1996; Turk 1996; Lolic et al. 2010; Mathioudakis et al. 2010; Borisova 2005; Ulubas and Ertunc 2005; Rana et al. 2007, 2009; Hassan et al. 2008; Keshavarz et al. 2009; Keshavarz and Shams-Bakhsh 2015; Pupola et al. 2011; Yaegashi et al. 2011; Kumar et al. 2012; Watpade et al. 2012; Brakta et al. 2013; Borisova et al. 2014; Cao et al. 2015; Han et al. 2015; Alemzadeh et al. 2016; Grimova et al. 2016b; Lemma et al. 2016; Winkowska et al. 2016). Symptoms and host(s) The virus-infected apple plants exhibit chlorotic spots with asymmetric leaf distortion, irregular diffused chlorotic rings, line patterns, dark green mottling, stem pitting, bark splitting, reduced size leaves, stunting, terminal dieback of some clones, and also fruit necrosis (Cropley 1964; Yanase 1974). Symptoms vary seasonally (Chairez and Lister 1973). The natural host range of this virus includes pear, peach, plum, cherry, and apricot (Nemeth 1986; Desvignes and Boye 1989). Transmission No natural vector is reported for this virus. The virus is transmissible by mechanical inoculation to a number of herbaceous hosts (Chenopodium and Nicotiana spp.). The use of infected clonal root stocks and use of infected trees as source of budwood propagation materials are the major sources of virus spread (Mink and Shay 1962; Luckwill and Campbell 1963; Zein and Zawarn 2005). The virus is transmissible by grafting (possibly by root grafts; Gilmer et al. 1971) but not by seed or pollen (Yoshikawa 2001). Virion properties and genome The virions are flexuous filaments, 680–780 nm long and 12 nm wide (Bar-Joseph et al. 1974), with distinct cross banding, made up of coat protein subunits of a single type, encapsidating a singlestranded, positive-sense RNA of about 7,555 nt, excluding the poly-A tail (M58152 = NC_001409) (Yoshikawa and Takahashi 1988; Sato et al. 1993; Zein and Zawarn 2005). The virus genome possesses three slightly overlapping open reading frames (ORFs) encoding proteins with approximate molecular weights of 216.5, 50 and 22 kDa, respectively. The ORF1 codes a 216.5 kDa protein containing the consensus motifs of methyltransferase, papain-like protease, nucleotide triphosphate-binding helicase, and RNA polymerase (German et al. 1990). The ORF2 (50 kDa) codes for the putative movement protein, and the ORF3 (21–22 kDa) codes for the coat protein (German et al. 1990). The largest protein (216.5 kDa) is directly expressed from the genomic RNA, while the other two are expressed via subgenomic mRNAs (German et al. 1992; Candresse et al. 1996; Adams et al. 2004; Zheng et al. 2007; Dhir et al. 2013).

Apple dimple fruit viroid Taxonomic position Genus: Apscaviroid

(ADFVd)

Family: Pospiviroidae

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Malus domestica (Apple)

Geographical distribution ADFVd was first reported from Italy in commercial trees of the apple cultivars “Starking Delicious,” “Royal Gala,” “Golden Delicious,” “Annurca,” and “Red Delicious” (Di Serio et al. 1996, 1998, 2001, 2002, 2003). The agent was then reported from “Starking Delicious” in Lebanon (Afechtal et al. 2007; Choueiri et al. 2007), from “Fuji,” and “Jonagold” in Japan (He et al. 2010; Kasai et al. 2017), and from “Fuji” and “Gala” in China (Ye et al. 2013). Symptoms and host(s) Typical symptoms observed in Italy on apple fruits of cv. Starking Delicious were depressed green spots 3–4 mm in diameter scattered over the whole fruit surface and around the calyx, where they sometimes merged to give large discolored areas (Di Serio et al. 1998, 2000; Ye et al. 2013). On apples in Japan, the leaves and branches of the infected trees did not have any disease symptoms, but the fruits were dimpled and yellow. The infected scions were top-grafted onto a healthy “Fuji” apple tree, and yellow dimple fruits were reproduced in the second growing season (He et al. 2010). Transmission ADFVd can be transmissible by grafting and also by mechanical stem slashing to young apple seedlings, using purified circular RNA. ADFVd has a narrow host range, but it was shown that the viroid can be transmitted artificially to a pear indicator (Di Serio et al. 2001). Virus-infected planting material is the main source of spread of this viroid (Malfitano et al. 2004). Etiology and genome properties The genome of the original isolate from Italy consists of a single-stranded circular RNA of 306 nt (NC_003463) reported in Italy (Di Serio et al. 1996). An isolate from Japan consisted of 303 nt was distinct from Italian isolates by 47 mutations (38 substitutions, 6 deletions, and 3 insertions) with an 84.9% overall nucleotide sequence homology (He et al. 2010). The isolate from China consisted of 306~307 nt with only two nucleotide substitutions compared to Italian isolates (Gora-Sochacka 2004; Giguere et al. 2014; Ye et al. 2013).

Apple fruit crinkle viroid

(AFCVd)

Taxonomic position AFCVd is a tentative member of the genus Apscaviroid and family Pospiviroidae Geographical distribution AFCVd was first identified from plants of Malus domestica in Japan in 1976 (Koganezawa and Ito 2003). The viroid-infected apple trees are distributed widely in apple-growing areas in Japan, but the disease incidence is sporadic (Ito et al. 1993; Koganezawa 2001). Symptoms and host(s) Apple fruit crinkle disease was recognized in the mid-1970s as a graft-transmissible disorder. The major symptoms consist of crinkling and dappling of the surface of mature fruit, although the severity of crinkling varies among the commercial cultivars, e.g., “Ohrin” tends to exhibit the most serious fruit deformation accompanied with scattered brown necrotic areas in the fruit flesh, “Nero 26” shows blister bark symptoms, and “Starking Delicious” produces only graft-transmissible bark disorders resembling

Malus domestica (Apple)

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apple measles caused by manganese toxicity or boron deficiency, without showing fruit symptoms (Ito et al. 1993; Ito and Yoshida 1998). Distinct strains are also reported from cultivated hops (Humulus lupulus) and Japanese persimmon (Diospyros kaki). Transmission The viroid is easily transmissible by budding, grafting, and chip budding and was transmissible mechanically to apple seedlings by razor-slashing with a purified preparation (Koganezawa et al. 1989; Ito et al. 1993). No attempts have been reported to inoculate AFCVd to woody hosts other than Malus species. Herbaceous hosts susceptible to AFCVd has not been reported so far, although cultivated hops are identified as a natural host for the viroid (Sano et al. 2008). Etiology and genome properties The genome of AFCVd consists of 369–372 nt (NC_003777). AFCVd is now listed as a tentative species of the genus Apscaviroid because its overall sequence homology to AGVd (85–87%) is just at the border of the demarcation criteria for viroid species (King et al. 2012).

Apple latent spherical virus Taxonomic position Genus: Cheravirus

(ALSV)

Family: Secoviridae

Geographical distribution Natural infection of ALSV in trees of Malus domestica was reported from Morioka, Japan, and other Asian countries (Mink et al. 1971; Li et al. 2000; Yoshikawa et al. 2000; Nakamura et al. 2011). Symptoms and host(s) The virus-infected apple trees did not exhibit any obvious symptoms in most of the apple cultivars (Ito and Yoshida 1997; Koganezawa et al. 1985). In Japan, some apple cultivars show fruit russet ring symptoms. Transmission The viroid is mechanically sap-transmissible to other plant species including those in the families Chenopodiaceae, Cucurbitaceae, Leguminaceae, Rosaceae, and Solanaceae (Igarashi et al. 2009). The virus is graft-transmissible and transmissible through infected vegetative propagating material. ALSV is seed-transmitted at a rate of ca. 4.5% in apple (Nakamura et al. 2011). Virion properties and genome The virions are non-enveloped and 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). Virions separate as two components, M and B, which are thought to contain two molecules of RNA 2 and a single molecule of RNA 1, respectively (Li et al. 2000, 2004). RNA1 is 6812 nt (AB030940 = NC_003787) excluding the 30 poly (A) tail and has a single open reading frame (ORF) encoding a replication-associated protein of 243 K (Li et al. 2000). RNA2 is 3384 nt (AB030941 = NC_003788) excluding the 30 poly (A) tail and has a single ORF encoding a 42-kDa movement protein (MP) on the N-terminal side and 3 capsid proteins in the C-terminal region (Koganezawa et al. 1985; Chunjiang et al. 2000; Li et al. 2000; Yoshikawa et al. 2000, 2006; Le Gall et al. 2007; Yoshikawa 2011; Sanfacon 2015; Thompson et al. 2017).

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Malus domestica (Apple)

Apple mosaic virus

(ApMV)

Taxonomic position Genus: Ilarvirus

Family: Bromoviridae

Geographical distribution ApMV was first reported in trees of Malus domestica by Bradford and Joly (1933) in the USA and is probably distributed worldwide (Wood et al. 1975; Bremer 1985; Howell et al. 1990; Turk 1996; Lee et al. 1998; Ghanem and Ashour 2002; Ulubas and Ertunc 2003; Akbas and Ilhan 2005; Yardimci and Eryigit 2005; Dursunoglu and Ertunc 2008; Yardimci et al. 2008; Hassan et al. 2008; Rana et al. 2009; Thokchom et al. 2009; Lakshmi et al. 2011; Padder et al. 2011; Paunovic et al. 2011; Petrzik and Lenz 2011; Kumar et al. 2012; Robertson 2012; Korkmaz et al. 2013; Watpade et al. 2013; Ertunc et al. 2014; Al-Jebr et al. 2005; Valasevich et al. 2015; Grimova et al. 2016a, b; Winkowska et al. 2016). Symptoms and host(s) ApMV-infected apple trees develop pale to bright cream spots on spring leaves as they expand. These spots may become necrotic after exposure to summer sun and heat. Most commercial cultivars are affected but vary in severity of symptoms. “Golden Delicious” and “Jonathan” are severely affected, whereas “Winesap” and “Mclntosh” are only mildly affected. Except in severe cases, a crop can still be produced by infected trees; yield reductions vary from 0–50%. In some cultivars, bud set is severely affected. ApMV has a wide host range, including woody and herbaceous plants. It is capable of infecting over 65 species in 19 families, by either experimental or natural routes (Grimova et al. 2016a). Horticulturally important host plants infected by this virus include apple (Malus domestica), pear (Pyrus communis), apricot (Prunus armeniaca), peach (P. persica), cherry (P. avium), plum (P. domestica), almond (P. amygdalus), strawberry (Fragaria spp.), raspberry (Rubus idaeus), blackberry (R. occidentalis), red currant (R. rubrum), and hazelnut (Corylus avellana). Other susceptible species include hop (Humulus lupulus), roses (Rosa spp.), wooly blackberry (Rubus canescens), R. ursinus, and further Prunus species such as P. cerasifera, P. instititia, P. mahaleb, P. salicina, P. serulata, P. triloba, P. cerasus, and blackthorn (P. spinosa) (Grimova et al. 2016a). ApMV has also been detected in silver birch (Betula pendula), white birch (B. papyrifiera), yellow birch (B. alleghaniensis), European mountain ash (Sorbus aucuparia), horse chestnut (Aesculus hippocastanum), red horse chestnut (A.  carnea), sweet buckeye (A. parviflora), bottle brush buckeye (A. flava), and hawthorn (Crataegus spp.) (Grimova et al. 2016a). ApMV has been detected in weeds that are often found in ApMV-infected hazelnut orchards, including shepherd’s needle (Scandix spp.), mugwort (Artemisia vulgaris), bellflower (Campanula spp.), wild clary (Salvia verbenaca), hemp nettle (Galeopsis spp.), self-heal (Prunella spp.), old man’s beard (Clematis vitalba), and lichens (Grimova et al. 2016a). Transmission There is no insect vector known for this virus. The virus is transmissible by mechanical sap-inoculation, and nearly 65 herbaceous plant species in 19 families are susceptible (EPPO 2000). The virus is transmissible by a vegetative propagation of root stocks and planting material from infected mother trees, and by graft-inoculation to woody plants. Root grafts appear to be the most important means of inter-tree transmission (Dhingra 1972). The virus is transmitted most probably by pollen in the same manner as other ilarviruses (thrips feeding) but not through seed.

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Virion properties and genome The virions are isometric and non-enveloped, with pleomorphic particles of 25 and 29 nm in diameter. The genome consists of three segmented ssRNAs: RNA1 consists of 3476 nt (AF174584 = NC _003464), RNA2 is 2979 nt (AF174585 = NC_003465), and RNA3 is 2056 nt (U15608 = NC_003480) (Alrefai et al. 1994, 1995; Sanchez-Navarro and Pallas 1994; Shiel et al. 1995; Shiel and Berger 2000; Petrzik and Lenz 2002; Thokchom et al. 2009; Scott 2011a, b).

Apple scar skin viroid Taxonomic position Genus: Apscaviroid

(ASSVd)

Family: Pospiviroidae

Geographical distribution ASSVd infection in trees of Malus domestica is widespread in Japan and China. The viroid is also reported from India, Iran, Australia, Argentina, Canada, France, Italy, Korea, and the USA (Agrios 1989; Hadidi et al. 1990; Zhu et al. 1995; Behl et al. 1998; Handa et al. 1998; Puchta et al. 1990; Desvignes et al. 1999; Koganezawa 2001; Lee et al. 2001; Koganezawa et al. 2003; Rana et al. 2009; Walia et al. 2009; Kaponi et al. 2010; Kumar et al. 2012; Nome et al. 2012; Yazarlou et al. 2012).

Symptoms and host(s) A wide range of malus species and apple varieties respond with scar skin, dappling, dimpling, cracking, rusty skin, and abscence of regular fruit pigmentation. In ASSVd-affected apple trees, the symptoms have been reported primarily on fruits. Early symptoms occur near the calyx end of the fruit as small reddish brown areas consisting of tiny raised structures. These areas coalesce to form reddish brown patches containing small areas of brownish scarlike tissues (Baumann and Engel 1972). The symptoms appear on a large area of the fruit surface but, especially near the calyx end. Two or 3 weeks later, russeted and scarred brown areas with numerous tiny brown fissures appear. Some fissures develop into deep cracks. The diseased fruits are only onethird of normal size and are unmarketable. Known hosts are confined to Malus and Pyrus in the Rosaceae. The viroid causes dimple fruit in Japanese pear (Pyrus pyrifolia), but there are no symptoms in the other pears (Pyrus bretschneideri, P. serotina, P. ussuriensis, and P. communis) (Desvignes et al. 1999).

Transmission There is no evidence for transmission of the virus by any vector. The viroid is mechanically saptransmissible; and experimentally by grafting and budding or by a razor-slash method. The field spread is believed to result from root contacts. This viroid is not seed-transmitted (Howell et al. 1998). The primary spread of this virod disease takes place through the use of infected planting material.

Virion properties and genome The genome consists of a single-stranded circular RNA of 329 nt, which is noncoding (X17696 = NC_001340) (Hashimoto and Koganezawa 1987; Koganezawa 1988; Puchta et al. 1990; Gora-Sochacka 2004; Walia et al. 2009; Flores et al. 2011; Yazarlou et al. 2012; Giguere et al. 2014).

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Malus domestica (Apple)

Apple stem grooving virus Taxonomic position Genus: Capillovirus

(ASGV)

Family: Betaflexiviridae

Geographical distribution ASGV infection is distributed worldwide wherever trees of Malus domestica is cultivated, such as North America, South America, Australia, China, India, Italy, Japan, the Netherlands, New Zealand, Portugal, the Czech Republic, Tunisia, Ethiopia, Brazil, Egypt, Korea and the UK (Uyemoto and Gilmer 1971; Plese et al. 1975; Fuchs 1981; Turk 1996; Kinard et al. 1996; Kummert et al. 1998; Polak and Zieglerova 2001; Caglayan et al. 2003; Kundu 2003; Al-Jebr et al. 2005; Hassan et al. 2008; Rana et al. 2009; Negi et al. 2010; Youssef et al. 2010; Massart et al. 2011; Pupola et al. 2011; Kumar et al. 2012; Brakta et al. 2013; Liu et al. 2013; Borisova et al. 2014; Han et al. 2015; Han et al. 2016; Grimova et al. 2016b; Lemma et al. 2016; Winkowska et al. 2016; Skhiri et al. 2017; Souza et al. 2017). Symptoms and host(s) The virus is not known to induce obvious symptoms in most commercial cultivars. Virginia crab develops long grooves on the woody stem, usually apparent only when the bark is removed. The grooves may be single and scattered or may be congregated in flame-like patterns. A swelling at the union with the root stock occurs frequently. Removal of bark from infected trees reveals a groove encircling the woody cylinder at the graft union that is matched by a ridge of necrotic phloem, and such grafts break easily (Waterworth and Uyemoto 1980; Ahman 1995). When the symptoms at the union are severe, the trees may be dwarfed or may decline. In the following season, leaves turn pale green and drop prematurely or may develop chlorotic flecks (Vander Meer 1976; Yanase 1983). The natural host range of ASGV includes Japanese apricot, cherry, citrus, European pear, kiwifruit, and lily. Transmission No vector has been identified for this virus. The virus is mechanically sap-transmissible. The virus is transmissible by grafting, and disseminated by infected propagative material (Yanase 1983). Virion properties and genome The virions are flexuous filaments and non-enveloped, with a clear modal length of 620 nm and 12 nm wide, encapsidating a positive-sense ssRNA genome of 6495 nt (D14995 = NC_001749) excluding the poly (A) tail at the 30 end of the genome. The coat protein subunit has a molecular mass of 27 kDa (Yoshikawa and Takahashi 1988; Yoshikawa et al. 1992; Terauchi et al. 1997; Nickel et al. 2001; Adams et al. 2004).

Apple stem pitting virus Taxonomic position Genus: Foveavirus

(ASPV)

Family: Betaflexiviridae

Geographical distribution The virus has been distributed worldwide wherever trees of Malus domestica are cultivated, including Australia, Austria, Bulgaria, Canada, China, the former Czechoslovakia, the Czech Republic, Denmark, Ethiopia, France, Germany, Greece, Hungary, India, Israel, Italy, Japan, Korea Republic, the

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Netherlands, New Zealand, Norway, Poland, Portugal, Romania, South Africa, Spain, Sweden, Switzerland, the UK, the USA, the former USSR, Uruguay, and the former Yugoslavia (Cropley 1968; Li and Mink 1989; Jelkmann 1997; Koganezawa and Yanase 1990; Caglayan et al. 2003; Kundu 2003; Hassan et al. 2008; Rana et al. 2009; Dhir et al. 2010; Lolic et al. 2010; Mathioudakis et al. 2010; Youssef et al. 2010; Jelkmann and Paunovic 2011; Li et al. 2011; Pupola et al. 2011; Kumar et al. 2012; Brakta et al. 2013; Yoon et al. 2014; Cao et al. 2015; Grimova et al. 2016b; Lemma et al. 2016; Winkowska et al. 2016; Hu et al. 2017).

Symptoms and host(s) The virus induces symptoms of epinasty and decline in the indicator Malus pumila “Spy 227”. Depending on the virus strain and apple cultivar, the symptoms are formation of distinct or coalescing pits in the xylem extending up to graft union. Although causing no marked symptoms in commercial apple cultivars, this virus may have a detrimental effect on the growth and cropping of some cultivars. Peg-like protrusions on the inner bark surface match the pitlike indentations on the opposite surface of the wood. Infected Virginia crab trees produce a characteristic fruit symptom called flute fruit observed as an abnormal fruit length compression and furrows running from the stem to calyx ends producing a fluted appearance. The natural hosts of this virus are Malus domestica, M. sieboldii, Pyrus communis, and Cydonia oblonga.

Transmission The virus is transmitted by means not involving a vector. The virus is transmissible by mechanical sapinoculation, and by grafting. The virus is not transmissible by contact between plants nor by seed and pollen. The use of bud sticks from infected plants is the major mode of spread of this disease.

Virion properties and genome The virions are flexuous filaments and non-enveloped, with a clear modal length of 800 nm and 12–15 nm wide. The genome is a single molecule of single-stranded, positive-sense RNA of 9332 nt (D21829 = NC_003462) (Jelkmann 1994) and comprises five ORFs. Analysis of the five ORFs shows the positive strand encoding for proteins with molecular masses of 247 k (ORF1), 25 k (ORF2), 13 k (ORF3), 7–8 k (ORF4), and 42–44 k (ORF5) (Koganezawa and Yanase 1990; Yoshikawa et al. 2001; Adams et al. 2004; Liu et al. 2012).

Cherry leaf roll virus Taxonomic position Genus: Nepovirus

(CLRV)

Family: Secoviridae

CLRV infection in trees of Malus domestica was reported from New Zealand by Woo et al. (2012). The virus-infected apple trees exhibit symptoms of necrotic spots, mottled lesions and leaf tip deformation symptoms. The virus is transmitted by nematode vectors in a non-persistent manner. The virus is also transmissible by mechanical sap-inoculation to a wide range of herbaceous species and by grafting. Vegetative propagation of virus-infected planting material is an efficient mode of spread. For more details of CLRV, refer to Prunus avium.

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Malus domestica (Apple)

Cherry rasp leaf virus Taxonomic position Genus: Cheravirus

(CRLV)

Family: Secoviridae

CRLV infection in trees of Malus domestica was reported from the USA, Canada, and Iran (James et al. 2001; James and Upton 2002; Ahmad 2016). The virus-infected apple trees produce small flattened fruit, reduced lateral branch growth, and upward rolling leaves. The virus is transmitted by nematode vectors Xiphinema americanum and X. californicum (Brown and Trudgill 1998). The virus is transmissible by mechanical sap-inoculation, and also through budding and grafting. For more details of CRLV, refer to Prunus avium.

Cucumber mosaic virus

(CMV)

Taxonomic position Genus: Cucumovirus

Family: Bromoviridae

CMV infection in trees of Malus domestica was reported from China (Hu et al. 2016a). The virus-infected apple plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Hop stunt viroid

(HpSVd)

Taxonomic position Genus: Hostuviroid

Family: Pospiviroidae

HpSVd infection in plants of Malus domestica was reported from Greece (Kaponi et al. 2010). The viroid is mechanically sap-transmissible. Propagation with shoots from infected plants is the primary cause of this viroid spread. For more details of HpSVd, refer to Humulus lupulus.

Peach latent mosaic viroid Taxonomic position Genus: Pelamoviroid

(PLMVd)

Family: Avsunviroidae

PLMVd infection in trees of Malus domestica was reported from Egypt (El-Dougdoug 1998). The viroid-infected apple plants exhibit a canker on bark, and scab, and streaking on the fruit. The viroid is transmissible by mechanical sap-inoculation, and also by grafting and budding. The use of viroidinfected vegetative planting material is one of the causes for disease spread. For more details of PLMVd, refer to Prunus persica.

Pear blister canker viroid Taxonomic position Genus: Apscaviroid

(PBCVd)

Family: Pospiviroidae

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PBCVd infection in plants of Malus domestica was reported from Greece (Kaponi et al. 2010). The viroid can be transmitted by grafting and budding, and mechanical inoculation. For more details of PBCVd, refer to Pyrus communis.

Prunus necrotic ringspot virus Taxonomic position Genus: Ilarvirus

(PNRSV)

Family: Bromoviridae

PNRSV infection in trees of Malus domestica was reported from India and China (Chandel et al. 2008; Rana et al. 2009; Hu et al. 2016b). The virus-infected apple plants exhibit necrotic symptoms on the leaves. The virus is present on pollen, entering the host through injuries caused by thrips while feeding. The virus is transmissible by mechanical sap-inoculation, and by grafting. The use of bud sticks from virus-infected plants for further propagation of planting material is the major cause of virus spread. The virus is not transmissible by contact between plants. For more details of PNRSV, refer to Prunus persica.

Tobacco mosaic virus

(TMV)

Taxonomic position Genus: Tobamovirus

Family: Virgaviridae

TMV was isolated from trees of Malus domestica in France (Kirkpatrick and Lindner 1964; Lemoine and Morand 1993). There is no known vector for this virus. The virus is transmissible by mechanical sap-inoculation, and also by contact between plants. The virus is transmissible by grafting and is seedtransmitted in Malus domestica up to 35% (Gilmer and Wilks 1967). For more details of TMV, refer to Nicotiana tabacum.

Tobacco ring spot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

TRSV infection in trees of Malus domestica was reported from Canada (Lana et al. 1983). The virusinfected apple plants exhibit stem pitting, necrosis, and breaking or separation of scion/rootstock at the graft union. Foliage was sparse and leaves were chlorotic and diffusely mottled. The virus is transmitted by nematode vectors, Xiphinema americanum and X. californicum, in a non-persistent manner, and also by mechanical sap-inoculation (Brown and Trudgill 1998). The virus is not transmissible by contact between plants. For more details of TRSV, refer to Nicotiana tabacum.

Tomato bushy stunt virus Taxonomic position Genus: Tombusvirus

(TBSV)

Family: Tombusviridae

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Malus domestica (Apple)

TBSV infection in trees of Malus domestica was reported from Canada (Allen 1969). The virus is transmissible by mechanical sap-inoculation, and also by grafting. The virus is seed-transmissible in apple. The virus is not transmissible by contact between plants. There is no known vector for this virus (Allen 1969). For more details of TBSV, refer to Solanum lycopersicum.

Tomato ringspot virus Taxonomic position Genus: Nepovirus

(ToRSV)

Family: Secoviridae

ToRSV is distributed in most parts of the world wherever trees of Malus domestica are grown (Stouffer and Uyemoto 1976; Stouffer et al. 1977; Lister et al. 1980; Parish and Converse 1981; Rosenberger et al. 1983; Bitterlin et al. 1984; Moini 2010). This disease is also known as apple union necrosis and decline in apple wherein the bark of affected trees becomes thickened and disorganized at the graft union, and a row of pits and grooves occurs along the graft union. Brown, necrotic tissues extend from these pits across the union, weakening it so that affected trees break at the union. The disease is most severe on Delicious and some other cultivars when propagated on MM 106 (EPPO 1991). The leaves are sparse and small and have a pale, dull green color on virus-infected apple trees. The terminal growth is reduced and the twigs express some rosetting. Diseased trees tend to flower heavily and commonly set a large number of small fruit with a brightly colored blush. The lateral leaf and flower buds may be killed, and the internodal length of the shoots is reduced. In severely affected trees, there is a proliferation of rootstock sprouts, and breakage at the union is not uncommon. In Iran, ToRSVaffected apple trees have shown yellowing of main veins, mosaic, and necrotic lesions (Moini 2010). The virus is transmitted by dagger nematodes, Xiphinema americanum, X. bricolense, and X. californicum, in a non-persistent manner (Forer et al. 1984; Georgi 1988; Brown and Trudgill 1998). Mechanical inoculation of ELISA-positive samples to Chenopodium amaranticolor-induced local lesions and on Nicotiana tabacum cv. Samson resulted in systemic infection. Local lesions were observed on N. rustica and chlorotic local lesions on C. quinoa; infection of Cucumis sativus remained symptomless. The use of vegetative propagative material from virus-infected plants is an efficient mode of spread (Forer et al. 1984). For more details of ToRSV, refer to Solanum lycopersicum.

Tulare apple mosaic virus Taxonomic position Genus: Ilarvirus

(TAMV)

Family: Bromoviridae

Geographical distribution TAMV was originally identified in trees of Malus domestica in Tulare County, California, USA (Yarwood 1955). Natural infection of the virus has only been detected in California and France (Brunt et al. 1996). Symptoms and host(s) The leaves of virus-infected apple trees develop severe symptoms of mosaic, chlorosis, vein-clearing, and ring-spotting. Patterns often appear on veins and adjacent tissues, and spots are not sharply delineated but are spread irregularly along the veins. Fruits develop no diagnostic symptoms in most apple cultivars tested (WSU 2003).

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Transmission The virus is mechanically sap-transmissible to several (three to nine) families of herbaceous plants (Mink and Bancroft 1962). Vegetative propagation of virus-infected planting material is an efficient mode of spread. The virus is not transmitted by seed. Virion properties and genome The virions are isometric and 33 nm in diameter, and the genome is a ssRNA (Fulton 1971; Lister and Saksena 1976). The genome consists of three segments of linear, positive-sense, ssRNA: RNA1 consists of 3,459 nt (AF226160 = NC_003833), RNA2 is 2944 nt (AF226161 = NC_003834), and RNA3 is 2,229 nt (AF226162 = NC_003835) respectively (Scott et al. 2003; Scott 2011a, b).

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Afechtal M, Choueiri E, El-Zammar S, Jreijiri F, Hobeika C, Myrta A, Di Serio F (2007) Identification and characterization of Apple dimple fruit viroid in Lebanon. J Plant Pathol 89(3):S27 Agrios GN (1989) Apple scar skin. In: Fridlund PR (ed) Virus and viruslike diseases of pome fruits and simulating noninfectious disorders. Washington State University, Pullman, pp 63–68 Ahmad C (2016) Cherry rasp leaf virus. A guide for diagnosis and detection of quarantine pests. Islamic Republic of Iran, Ministry of Jihad-e-Agriculture, Plant Protection Organization, p 11 Ahman G (1995) Viruses in apple. Vaxtskyddsnotiser 59:52–55 Akbas B, Ilhan D (2005) Wide spread distribution of Apple mosaic virus on apple in Turkey. Plant Dis 89:1010 Alemzadeh E, Katsiani AT, Efthimiou K, Katis NI (2016) Occurrence of Apple chlorotic leaf spot virus in apple and quince in Southern Iran. J Plant Pathol 98:177 Al-Jebr K, Ismaeil F, Mando MJ, Al-Saadoun E, Al-Chaabi S (2005) First record of pome fruit viruses in Syria. J Plant Pathol 87(3):243 Allen WR (1969) Occurrence and seed transmission of Tomato bushy stunt virus in apple. Can J Plant Sci 49:797–799 Alrefai RH, Shiel PJ, Domier LL, D’Arcy CJ, Berger PH, Korban SS (1994) The nucleotide sequence of Apple mosaic virus coat protein gene has no similarity with other Bromoviridae coat protein genes. J Gen Virol 75:2847–2850 Alrefai RH, Domier LL, Korban SS (1995) The complete nucleotide sequence of apple mosaic virus RNA-3. Arch Virol 140:1247–1256 Barba M, Clark MF (1986) Detection of strains of Apple chlorotic leaf spot virus by F(ab)2-based indirect ELISA. Acta Hortic 193:297–304 Bar-Joseph M, Hull MR, Lane LC (1974) Biophysical and biochemical characterization of Apple chlorotic leaf spot virus. Virology 62:563–566 Baumann G, Engel G (1972) A graft transmissible disease of golden delicious. Erwerbsobstbau 14:58–59 Behl MK, Khurana SMP, Parakh DB (1998) Bumpy fruit and other viroid and viroid-like diseases of apple in HP, India. Acta Hortic 472:627–629 Bitterlin MW, Gonsalves D, Cummins JN (1984) Irregular distribution of Tomato ringspot virus in apple trees. Plant Dis 68:567–571 Borisova A (2005) Preliminary results of the study on the spread of Apple chlorotic leaf spot virus (ACLSV) in different fruit tree species in Kyustendil region of Bulgaria. Not Bot Hortic Agrobot Cluj 33:81–85 Borisova A, Borovinova M, Kamenova I (2014) Major diseases of apple trees in Kyustendil region of Bulgaria. Turkish J Agric Nat Sci Spec Issue 1:695–700 Bradford FC, Joly L (1933) Infectious variegation in the apple. J Agric Res 46:901–908 Brakta A, Thakur PD, Handa A (2013) First report of apple top working disease caused by viruses (apple stem grooving virus, apple chlorotic leaf spot virus, and apple stem pitting virus) in apple in India. Plant Dis 97:1001 Bremer K (1985) Hedelma-ja marjakasvien taudit. Kasvinsuojeluseuran julkaisu no. 76, 75 p Brown DJF, Trudgill DL (1998) Nematode transmission of plant viruses: a 30 year perspective. Host pathogen interactions and crop protection. SCRI Annual Report, pp 121–125 Brunt AA, Crabtree K, Dallwitz MJ, Gibbs AJ, Watson L (eds) (1996) Viruses of plants. Descriptions and lists from the VIDE database. CAB International, Wallingford, 1484 pp

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Waterworth HE, Uyemoto JK (1980) Symptoms incited by apple type II virus isolates in the Virginia crab apple trees. Plant Dis 64:562–563 Watpade S, Raigond B, Thakur PD, Handa A, Pramanick KK, Sharma YP, Tomar M (2012) Molecular detection of latent Apple chlorotic leaf spot virus in elite mother plants of apple. Indian Journal of Virology 23(3): 359–363 Watpade S, Raigond B, Pramanick KK, Sharma N, Handa A, Sharma U (2013) Simultaneous detection of Apple chlorotic leaf spot virus and Apple mosaic virus in crab apples and apple rootstocks by duplex RT-PCR. Scientia Horticulturae 164: 88–93 Winkowska L, Grimova L, Rysanek P (2016) Quantitative detection of four pome fruit viruses in apple trees throughout the year. Phytopathol Mediterr 55:207–224 Woo ENY, Clover GRG, Pearson MN (2012) First report of Cherry leaf roll virus (CLRV) in Malus domestica. Australas Plant Dis Notes 7:151–156 Wood GA, Chamberlain EE, Atkinson JD, Hunter JA (1975) Field studies with Apple mosaic virus. N Z J Agric Res 18:399–404 WSU (2003) “Virus free” stone & pome fruit – pome fruit diseases caused by viruses (NRSP5) 2001. Washington State University. National Research Support Project 5. Supporting Research & Development. Pome Fruit-Virus Detection Procedures (NRSP5, Indexing Procedures); at http://nrsp5.prosser.wsu.edu/index.html Yaegashi H, Yoshikawa N, Candresse T (2011) Apple chlorotic leaf spot virus in pome fruits. In: Hadidi A, Barba M, Candresse T, Jelkmann W (eds) Virus and virus-like diseases of pome and stone fruits. APS Press, St. Paul, pp 17–22 Yanase H (1974) Studies on apple latent viruses in Japan: The association of apple topworking with apple latent viruses. Bull Fruit Tree Res Stn Jpn Ser C (Morioka) 1:47–109 Yanase H (1983) Back transmission of apple stem grooving virus to apple seedlings and induction of symptoms of apple topworking disease in Mitsuba Kaido (Malus sieboldii) and Kabano Zumi (Malus sieboldii var. arborescens) rootstocks. Acta Hortic 130:117–122 Yardimci N, Eryigit H (2005) New record of apple mosaic virus on apple cultivars in south-west Turkey. Australas Plant Pathol 34:603 Yardimci N, Cevik B, Eryigit H (2008) Detection of Apple mosaic virus on apple cultivars growing in south-west Turkey by ELISA and RT-PCR methods. Acta Hortic (ISHS) 781:561–565 Yarwood CE (1955) Mechanical transmission of an Apple mosaic virus. Hilgardia 23:613–628 Yazarlou A, Jafarpour B, Habili N, Randles J (2012) First detection and molecular characterization of new Apple scar skin viroid variants from apple and pear in Iran. Australas Plant Dis Notes 7:99–102 Ye T, Chen SY, Wang R, Hao L, Chen H, Wang N, Guo LY, Fan ZF, Li SF, Zhou T (2013) Identification and molecular characterization of Apple dimple fruit viroid in China. J Plant Pathol 95:637–641 Yoon JY, Joa JH, Choi KS, Do KS, Lim HC, Chung BN (2014) Genetic diversity of a natural population of Apple stem pitting virus isolated from apple in Korea. Plant Pathol J 30:195–199 Yoshikawa N (2001) Apple chlorotic leaf spot virus. CMI AAB Description of Plant Viruses, no.386 (No. 30 revised) Yoshikawa N (2011) Cheravirus. Sequiviridae. In: The Springer Index of Viruses. Springer, New York, pp 1763–1768. https://doi.org/10.1007/978-0-387-95919-1_286 Yoshikawa N, Takahashi T (1988) Properties of RNAs and proteins of apple stem grooving and apple chlorotic leaf spot viruses. J Gen Virol 69:241–245 Yoshikawa N, Sasaki E, Kato M, Takahashi T (1992) The nucleotide sequence of Apple stem grooving capillovirus genome. Virology 191(1):98–105 Yoshikawa N, Takahashi T, Ito T, Yoshida K, Koganezawa H (2000) Nucleotide sequence and genome organization of apple latent spherical virus: a new virus classified into the family Comoviridae. J Gen Virol 81:541–547 Yoshikawa N, Matsuda H, Oda Y, Isogai M, Takahashi T, Ito T, Yoshida K (2001) Genome heterogeneity of Apple stem pitting virus in apple trees. Acta Hortic 550:285–290 Yoshikawa N, Okada K, Asanuma K, Watanabe K, Igarashi A, Li C, Isogai M (2006) A movement protein and three capsid proteins are all necessary for the cell-to-cell movement of Apple latent spherical cheravirus. Arch Virol 151:837–848 Youssef SA, Moawad SM, Nosseir FM, Shalaby AA (2010) Detection and identification of Apple stem pitting virus and Apple stem grooving virus affecting apple and pear trees in Egypt. 21st International Conference on Virus and other Graft Transmissible Diseases of Fruit Crops. Julius-Kühn-Archiv 427: 248–252 Zein SN, Zawarn HS (2005) Apple chlorotic leaf spot trichovirus. Int J Virol 1:19–19 Zheng YY, Wang GP, Hong N, Song YS, You H (2007) Partial molecular characterization of Apple chlorotic leaf spot virus from peach and apple trees and prokaryotic expression for cp gene. Acta Phytopathol Sin 37:356–361 Zhu SF, Hadidi A, Hammond RW, Yang X, Hansen AJ (1995) Nucleotide sequence and secondary structure of pome fruit viroids from dapple apple diseased apples, pear rustskin diseased pears and apple scar skin symptomless pears. Acta Hortic 386:554–559

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Malus sylvestris (European crab apple)

Malus sylvestris (European crab apple) Family: Rosaceae

Edible fruit

Apple chlorotic leaf spot virus Taxonomic position Genus: Trichovirus

(ACLSV)

Family: Betaflexiviridae

ACLSV infection in plants of Malus sylvestris was reported from China (Li et al. 2017). No vector is reported for this virus. The virus is mechanically sap-transmissible and also transmissible through grafting. The use of infected propagation material is regarded as the main means of dissemination. For more details of ACLSV, refer to Malus domestica.

Apple stem grooving virus Taxonomic position Genus: Capillovirus

(ASGV)

Family: Betaflexiviridae

ASGV infection in plants of Malus sylvestris was reported from the USA, China, and the UK (Lister et al. 1965; De Sequeira 1967; Li et al. 2017). The virus generally causes latent infection in most of the commercial cultivars of European crab apple. The virus produces stem grooving, swelling of graft unions, and symptoms of graft-union-necrosis in Virginia Crab. No vector has been identified for this virus. The virus is mechanically sap-transmissible. The virus is also transmissible by grafting and is disseminated by infected propagative material. For more details of ASGV, refer to Malus domestica.

Apple stem pitting virus Taxonomic position Genus: Foveavirus

(ASPV)

Family: Betaflexiviridae

ASPV infection in plants of Malus sylvestris was first reported from the USA (Smith 1954). The virusinfected European crab apple plants exhibit symptoms of stem pitting and epinasty. The virus is transmitted by means not involving a vector. Virus is transmissible by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants. Use of bud sticks from infected plants is the major mode of spread of this disease. For more details of ASPV, refer to Malus domestica.

Prunus necrotic ringspot virus Taxonomic position Genus: Ilarvirus

(PNRSV)

Family: Bromoviridae

PNRSV infection in plants of Malus sylvestris was reported from China (Li et al. 2017). The virus is present on pollen, entering the host through injuries caused by thrips while feeding. The virus is

Malva neglecta (Buttonweed)

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transmissible by mechanical sap-inoculation, and by grafting. The use of bud sticks from virus-infected plants for further propagation of planting material is the major cause of virus spread. The virus is not transmissible by contact between plants. For more details of PNRSV, refer to Prunus persica.

Tulare apple mosaic virus Taxonomic position Genus: Ilarvirus

(TAMV)

Family: Bromoviridae

TAMV infection in plants of Malus sylvestris was first reported from California, USA by Yarwood (1955). This virus is also reported from France and California (USA). Virus-infected European crab apple plants exhibit conspicuous leaf chlorosis symptoms. The virus is mechanically sap-transmissible, and the inoculated apple trees exhibited mild to severe symptoms of mosaic, chlorosis, vein-clearing, and ringspotting. The use of infected vegetative planting material is the primary source of virus spread. The virus is graft-transmitted but is not transmitted by seed. For more details of TAMV, refer to Malus domestica.

References De Sequeira OA (1967) Studies on a virus causing stem grooving and graft-union abnormalities in Virginia Crab apple. Ann Appl Bio 60:59–66 Li Y, Deng C, Bian Y, Zhao X, Zhou Q (2017) Characterization of apple stem grooving virus and apple chlorotic leaf spot virus identified in a crab apple tree. Arch Virol 162:1093–1097 Lister RM, Bancroft JB, Nadakavukaren MJ (1965) Some sap-transmissible viruses from apple. Phytopathology 55:859 Smith WW (1954) Occurrence of “stem pitting” and necrosis in some body stocks of apple trees. Proc Am Soc Hortic Sci 63:101–113 Yarwood CE (1955) Mechanical transmission of an Apple mosaic virus. Hilgardia 23:613–628

Malva neglecta (Buttonweed) Family: Malvaceae

Malva mosaic virus Taxonomic position Genus: Potexvirus

Weed host

(MalMV)

Family: Alphaflexiviridae

Geographical distribution MalMV infection in plants of Malva neglecta was reported from Canada (Cote et al. 2008). Symptoms and host(s) The virus-infected button weed plants exhibit mosaic symptoms. Transmission The virus is transmissible by mechanical inoculation but not by vectors.

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Malva parviflora (Cheese weed mallow)

Virion properties and genome The virions are flexuous filaments and 470–580 nm in length and 13 nm in diameter. The genome consists of a single linear molecule of positive-sense ssRNA of 6858 nt (DQ660333 = NC_008251) and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type and 18–27 kDa in size (Adams et al. 2004; Cote et al. 2008).

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Malva neglecta was reported from Italy (Grieco et al. 2000). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Malva neglecta was reported from Cyprus (Papayiannis et al. 2011). The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. The virus is transmitted by grafting but not by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Cote F, Pare C, Majeau N, Bolduc M, Leblanc E, Bergeron MG, Bernardy MG, Leclerc D (2008) Nucleotide sequence and phylogenetic analysis of a new potexvirus: Malva mosaic virus. Infect Genet Evol 8(1):83–93 Grieco PD, Conte D, Munno I, Nuzzaci M, de Stradis A (2000) Tomato spotted wilt virus (TSWV) on weeds and wild plants in Metapontino, Basilicata. Informatore Fitopatol 50:43–46 Papayiannis L, Katis NI, Idris AM, Brown JK (2011) Identification of weed hosts of Tomato yellow leaf curl virus in Cyprus. Plant Dis 95:120–125

Malva parviflora (Cheese weed mallow) Family: Malvaceae

Abutilon mosaic virus Taxonomic position Genus: Begomovirus

Weed host

(AbMV)

Family: Geminiviridae

Malva parviflora (Cheese weed mallow)

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AbMV infection in plants of Malva parviflora was reported from Europe (Jeske and Schuchalter-Eicke 1984; Abouzid and Jeske 1986; Wege et al. 2000). The virus-infected cheese weed mallow plants exhibit symptoms of yellow mosaic and stunting. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative non-propagative manner. The virus is mechanically sap-transmissible. For more details of AbMV, refer to Abutilon striatum (Syn.) Abutilon pictum.

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

(AMV)

Family: Bromoviridae

AMV infection in plants of Malva parviflora was reported from Saudi Arabia (Al-Shahwan et al. 2017). The virus is transmitted by several species of aphids in a non-persistent manner, and is also transmissible by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Beet curly top virus Taxonomic position Genus: Curtovirus

(BCTV)

Family: Geminiviridae

BCTV infection in plants of Malva parviflora was reported from California, USA (Creamer et al. 1996). The virus is transmitted by a leafhopper vector, Circulifer tenellus, in a persistent (circulative, nonpropagative) manner. The virus is not transmissible by mechanical inoculation. For more details of BCTV, refer to Beta vulgaris.

Cherry rasp leaf virus Taxonomic position Genus: Cheravirus

(CRLV)

Family: Secoviridae

CRLV infection in plants of Malva parviflora was reported from the USA (Villamor and Eastwell 2016). The virus-infected cheese weed mallow plants exhibit symptoms of chlorotic ring patterns, leaf blotch, and leaf deformations. The virus is transmitted by nematode vectors. The virus is mechanically sap-transmissible and also transmissible through budding and grafting. For more details of CRLV, refer to Prunus avium.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Malva parviflora was reported from New Zealand (Fletcher 2001). The virus-infected cheese weed mallow plants do not exhibit obvious symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

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Malva parviflora (Cheese weed mallow)

Cucumber vein yellowing virus Taxonomic position Genus: Ipomovirus

(CVYV)

Family: Potyviridae

CVYV infection in plants of Malva parviflora was reported from Spain (Janssen et al. 2002). The virusinfected cheese weed mallow plants exhibit vein-clearing and leaf chlorotic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CVYV, refer to Cucumis sativus.

Hollyhock leaf curl virus Taxonomic position Genus: Begomovirus

(HoLCV)

Family: Geminiviridae

HoLCV infection in plants of Malva parviflora was reported from Pakistan (Sattar et al. 2017). The virus-infected cheese weed mallow plants exhibit vein yellowing and chlorosis symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of HoLCV, refer to Alcea rosea (syn. Althaea rosea).

Lettuce mosaic virus Taxonomic position Genus: Potyvirus

(LMV)

Family: Potyviridae

LMV infection in plants of Malva parviflora was reported from California, USA (Zerbini et al. 1997). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of LMV, refer to Lactuca sativa.

Pepino mosaic virus Taxonomic position Genus: Potexvirus

(PepMV)

Family: Alphaflexiviridae

PepMV infection in plants of Malva parviflora was reported from Spain and Cyprus (Jorda et al. 2001a; Papayiannis et al. 2012). The virus is mechanically sap-transmissible. For more details of PepMV, refer to Solanum muricatum.

Squash leaf curl virus Taxonomic position Genus: Begomovirus

(SLCuV)

Family: Geminiviridae

Malva parviflora (Cheese weed mallow)

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SLCuV infection in Malva parviflora was reported from Jordan (Al-Musa et al. 2008). The virusinfected cheese weed mallow plants exhibit severe leaf curling, yellowing, and stunting of the whole plants. The whitefly Bemisia tabaci, transmits this virus in a circulative non-propagative manner. For more details of SLCuV, refer to Cucurbita pepo.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Malva parviflora was reported from Hawaii (Cho et al. 1986; Bautista et al. 1995). The virus is transmitted by the thrips vector, Frankliniella occidentalis, in a persistentpropagative manner, and is transmissible by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Malva parviflora was reported from Spain, Cyprus, and Israel (Cohen et al. 1988; Jorda et al. 2001b; Papayiannis et al. 2011). The virus-infected cheese weed mallow plants exhibit symptomless infections. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is transmissible by grafting but not by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

References Abouzid A, Jeske H (1986) The purification and characterization of gemini particles from Abutilon mosaic virus infected Malvaceae. J Phytopathol 115:344–353 Al-Musa A, Anfoka G, Misbeh S, Abhary M, Ahmad FH (2008) Detection and molecular characterization of Squash leaf curl virus (SLCV) in Jordan. J Phytopathol 156(5):311–316 Al-Shahwan IM, Abdalla OA, Al-Saleh MA, Amer MA (2017) Detection of new viruses in alfalfa, weeds and cultivated plants growing adjacent to alfalfa fields in Saudi Arabia. Saudi J Biol Sci 24(6):1336–1343 Bautista RC, Mau RFL, Cho JJ, Custer DM (1995) Potential of Tomato spotted wilt tospovirus plant hosts in Hawaii as virus reservoirs for transmission by Frankliniella occidentalis (Thysanoptera: Thripidae). Phytopathology 85:953–958 Cho JJ, Mau RFL, Gonsalves D, Mitchell WC (1986) Reservoir weed hosts of Tomato spotted wilt virus. Plant Dis 70:1014–1017 Cohen S, Kern J, Harpaz I, Ben-Joseph R (1988) Epidemiological studies of the tomato yellow leaf curl virus (TYLCV) in the Jordan Valley, Israel. Phytoparasitica 16:259–270 Creamer R, Lugue-Williams M, Howo M (1996) Epidemiology and incidence of Beet curly top geminivirus in naturally infected weed hosts. Plant Dis 80:533–535 Fletcher D (2001) New hosts of Alfalfa mosaic virus, Cucumber mosaic virus, Potato virus Y, Soybean dwarf virus, and Tomato spotted wilt virus in New Zealand. N Z J Crop Hortic Sci 29(3):213–217 Janssen D, Ruiz L, Velasco L, Segundo E, Cuadrado IM (2002) Non-cucurbitaceous weed species shown to be natural hosts of Cucumber vein yellowing virus in south eastern Spain. Plant Pathol 51:797 Jeske H, Schuchalter-Eicke G (1984) The Abutilon Mosaic Virus (AbMV) in different leaf tissues of several host species of Malvaceae. J Phytopathol 90:353–362

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Malva rotundifolia (Round-leaved Mallow)

Jorda C, Lazaro Perez A, Martinez-Culebras P, Lacasa A (2001a) First report of Pepino mosaic virus on natural hosts. Plant Dis 85:1292 Jorda C, Font I, Martinez P, Juarez M, Ortega A, Lacasa A (2001b) Current status and new natural hosts of Tomato yellow leaf curl virus (TYLCV) in Spain. Plant Dis 85:445 Papayiannis L, Katis NI, Idris AM, Brown JK (2011) Identification of weed hosts of Tomato yellow leaf curl virus in Cyprus. Plant Dis 95:120–125 Papayiannis LC, Kokkinos CD, Alfafo-Fernandez A (2012) Detection, characterization and host range studies of Pepino mosaic virus in Cyprus. Eur J Plant Pathol 132:1–7 Sattar MN, Qurashi F, Iqbal Z, Haider MS (2017) Molecular characterization of Hollyhock leaf curl virus and associated DNA-satellites infecting Malva parviflora in Pakistan. Can J Plant Pathol 39:229–234 Villamor DEV, Eastwell K (2016) Sambucus nigra subsp. Caerulea and Malva spp.: newly identified hosts of Cherry rasp leaf virus. Plant Dis 100:867 Wege C, Gotthardt RD, Frischmuth T, Jeske H (2000) Fulfilling Koch’s postulates for Abutilon mosaic virus. Arch Virol 145:2217–2225 Zerbini FM, Koike ST, Gilbertson RL (1997) Gazania spp.: a new host of lettuce mosaic potyvirus, and a potential inoculum source for recent lettuce mosaic outbreaks in the Salinas Valley of California. Plant Dis 81:641–646

Malva rotundifolia (Round-leaved Mallow) Family: Malvaceae

Medicinal

Tomato yellow leaf curl China virus Taxonomic position Genus: Begomovirus

(TYLCCNV)

Family: Geminiviridae

MaYLCV infection in plants of Malva rotundifolia was reported from China (Li et al. 2015). The virusinfected round-leaved mallow plants exhibit yellow vein and leaf crinkling symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of TYLCCNV, refer to Solanum lycopersicum.

References Li K, Zhang J, Jing CC, Wu GT, Sun XC, Qing L (2015) First report of Tomato yellow leaf curl China virus infecting Malva rotundifolia in China. J Plant Pathol 97:547

Malva sylvestris (Common mallow) Family: Malvaceae

Cherry rasp leaf virus Taxonomic position Genus: Cheravirus

Medicinal

(CRLV)

Family: Secoviridae

Malva sylvestris (Common mallow)

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CRLV infection in plants of Malva sylvestris was reported from the USA (Villamor and Eastwell 2016). The virus-infected common mallow plants exhibit symptoms of chlorotic ring patterns, leaf blotch, and leaf deformations. The virus is transmitted by nematode vectors. The virus is mechanically saptransmissible and also transmissible through budding and grafting. For more details of CRLV, refer to Prunus avium.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Malva sylvestris was reported from New Zealand and Hungary (Horvath et al. 1995; Fletcher 2001). The virus-infected common mallow plants do not exhibit obvious symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of CMV, refer to Cucumis sativus.

Malva vein clearing virus Taxonomic position Genus: Potyvirus

(MVCV)

Family: Potyviridae

Geographical distribution MVCV infection was first reported in plants of Malva sylvestris from Germany by Hein (1957). The virus spreads in Australia (Tasmania), Brazil, the former Czechoslovakia, Germany, Israel, Italy, Spain, Portugal, the USA (California), the former USSR, and the former Yugoslavia (Henriques and Henriques 1986; Lunello et al. 2009).

Symptoms and host(s) The virus-infected common mallow plants exhibit vein-clearing and yellow mosaic symptoms. Hollyhock is a natural host of this virus, and tomato has also been found infected.

Transmission The virus is transmitted by aphid vectors Aphis umbrella (syn. Aphis malvae Koch) and Myzus persicae in a non-persistent manner. The virus is also transmissible by mechanical inoculation to less than three plant families; only some genera of the Malvaceae are infected.

Virion properties and genome The virions are flexuous filaments and non-enveloped, with a clear modal length of 850–860 nm and 12 nm wide. The genome consists of a single molecule of linear, positive-sense ssRNA (Revers and Garcia 2015). Partial genome sequences of 1832 nt (FM212972) and 2078 nt (FJ561293) are available (Kitajima et al. 1962; Lunello et al. 2009; Wylie et al. 2017).

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Malva verticillata (Chinese mallow)

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Malva sylvestris was reported from Cyprus (Papayiannis et al. 2011). The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. The virus is transmissible by grafting but not by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

References Fletcher JD (2001) New hosts of Alfalfa mosaic virus, Cucumber mosaic virus, Potato virus Y, Soybean dwarf virus, and Tomato spotted wilt virus in New Zealand. N Z J Crop Hortic Sci 29:213–217 Hein A (1957) Beitrage zur Kenntnis der Viruskrankheiten an Unkrautern. I. Das Malva-Virus. Phytopathol Z 28:205–234 Henriques MIC, Henriques FS (1986) Association of malva vein-clearing virus and rhabdovirus like particles with mottle and vein clearing of malva plants. Can J Bot 64:85–89 Horvath J, Salamon P, Lesemann DE (1995) Natural occurrence of Cucumber mosaic cucumovirus on Malva sylvestris plant in Hungary. Acta Phytopathol Entomol 30:145–151 Kitajima EW, Costa AS, Carvalho AMB (1962) Particle morphology of the malva vein clearing virus. Bragantia 21. https://doi.org/10.1590/S0006-87051962000100073 Lunello P, Tourino A, Nunez Y, Ponz F, Sanchez F (2009) Genomic heterogeneity and host recovery of isolates of Malva vein clearing virus. Virus Res 140:91–97 Papayiannis L, Katis NI, Idris AM, Brown JK (2011) Identification of weed hosts of Tomato yellow leaf curl virus in Cyprus. Plant Dis 95:120–125 Revers F, Garcia JA (2015) Molecular biology of potyviruses. Adv Virus Res 92:101–199 Villamor DEV, Eastwell K (2016) Sambucus nigra subsp. Caerulea and Malva spp.: newly identified hosts of Cherry rasp leaf virus. Plant Dis 100:867 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354

Malva verticillata (Chinese mallow) Family: Malvaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Leafy vegetable

(CMV)

Family: Bromoviridae

CMV infection in plants of Malva verticillata was reported from China (Peng et al. 2015). The virusinfected Chinese mallow plants exhibit mosaic and stunting symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

References Peng H, Zhao C, Zhao X, Chen D, Sun X (2015) First report of Cucumber mosaic virus infecting Chinese mallow in China. J Phytopathol 163:1064–1068

Malvastrum spp.

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Malvastrum spp. Family: Malvaceae

Ageratum leaf curl virus Taxonomic position Genus: Begomovirus

Weed host

(ALCuV)

Family: Geminiviridae

ALCuV infection in plants of Malvastrum coromandelianum was reported from China (Huang and Zhou 2006). The virus-infected malvastrum plants exhibit leaf curling symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci, in a circulative non-propagative manner. For more details of ALCuV, refer to Ageratum spp.

Hollyhock leaf curl virus Taxonomic position Genus: Begomovirus

(HoLCV)

Family: Geminiviridae

EYVV infection in plants of Malvastrum coromandelianum was reported from Pakistan (Zia-UrRehman et al. 2018). The virus-infected malvastrum plants exhibit leaf curling symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci, in a circulative non-propagative manner. For more details of HoLCV, refer to Alcea rosea (syn. Althaea rosea).

Malvastrum bright yellow mosaic virus Taxonomic position Genus: Begomovirus

(MaBYMV)

Family: Geminiviridae

Geographical distribution MaBYMV infection in plants of Malvastrum spp. was reported from South Texas (USA) (Alabi et al. 2016). Symptoms and host(s) The virus-infected malvastrum plants exhibit bright yellow mosaic symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2609 nt (KU058856 = NC_031458) and DNA-B of 2604 nt (KU058860 = NC_031459) (Briddon 2001; Brown et al. 2015; Alabi et al. 2016; Zerbini et al. 2017).

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Malvastrum spp.

Malvastrum leaf curl Philippines virus Taxonomic position Genus: Begomovirus

(MaLCPHV)

Family: Geminiviridae

Geographical distribution MaLCPHV infection in plants of Malvastrum coromandelianum was reported from the Philippines (She and He, unpublished - KC577540). Symptoms and host(s) The virus-infected malvastrum plants exhibit leaf curl symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2742 nt (KC577540 = NC_021245) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). Malvastrum leaf curl Philippines deltasatellite DNA molecule is associated with MaLCPHV, and consists of 673 nt (KF433066) (Zhou 2013).

Malvastrum leaf curl virus Taxonomic position Genus: Begomovirus

(MaLCuV)

Family: Geminiviridae

Geographical distribution MaLCuV infection in plants of Malvastrum coromandelianum was reported from China (Huang and Zhou 2006). Symptoms and host(s) The virus-infected malvastrum plants exhibit leaf curl symptoms. The virus also naturally infects papaya causing leaf curl symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2745 nt (AJ971263 = NC_007724) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). Malvastrum leaf curl betasatellite DNA molecule is associated with MaLCuV, and consists of 1354 nt (AM072289) (Huang and Zhou 2006; Zhou 2013).

Malvastrum spp.

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Malvastrum yellow mosaic Helshire virus Taxonomic position Genus: Begomovirus

(MaYMHeV)

Family: Geminiviridae

Geographical distribution MaYMHeV infection in plants of Malvastrum americanum was reported from Jamaica and the Northern Caribbean (Graham et al. 2010). Symptoms and host(s) The virus-infected malvastrum plants exhibit foliar yellow mosaic symptoms. Transmission The transmission of MaYMHeV has not been investigated. It is likely that, in common with other begomoviruses, MaYMHeV will be transmitted by the whitefly Bemisia tabaci in a circulative, nonpropogative manner. Virion properties and genome The structure of MaYMHeV particles has not been investigated. In common with all geminiviruses the virions of MaYMHeV will likely be geminate (twinned icosahedra). The genome of MaYMHeV is poorly characterised. Only the sequence of a single cicular component, homologous to the genomes of monopartite begomoviruses and DNA A component of bipartite begomoviruses has been determined. The majority of begomoviruses native to the New World are bipartite, suggesting that the sequence of the genome of MaYMHeV is incomplete – missing the DNA B component. The genome contains a single molecule of circular single-stranded DNA (DNA-A) of 2609 nt (FJ600483) (Briddon 2001; Graham et al., 2010; Brown et al. 2015; Zerbini et al. 2017). The only available sequence of MaYMHeV encodes the five genes typically encoded by the DNA A components of bipartite begomoviruses or the genomes of monopartite begomoviruses native to the New World.

Malvastrum yellow mosaic Jamaica virus Taxonomic position Genus: Begomovirus

(MaYMJV)

Family: Geminiviridae

Geographical distribution MaYMJV infection in plants of Malvastrum americanum was reported from Jamaica and the Northern Caribbean (Graham et al. 2007, 2010). Symptoms and host(s) The virus-infected malvastrum plants exhibit yellow mosaic symptoms. Transmission The virus is transmitted by the B-biotype of the whitefly vector Bemisia tabaci, in a circulative nonpropagative manner.

M

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Malvastrum spp.

Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is a bipartite consisting of two circular, single-stranded DNA components. The DNA-A consists of 2612 nt (FJ601917), and DNA-B of 2585 nt (FJ600485) (Briddon 2001; Graham et al. 2010; Brown et al. 2015; Zerbini et al. 2017).

Malvastrum yellow mosaic virus Taxonomic position Genus: Begomovirus

(MaYMV)

Family: Geminiviridae

Geographical distribution MaYMV infection in plants of Malvastrum spp. was reported from China (Guo and Xie 2008; Guo et al. 2007). Symptoms and host(s) The virus-infected malvastrum plants exhibit yellow mosaic symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2728 nt (AM236755 = NC_008559) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017). A betasatellite of 1340 nt (AM236769) associated with the virus was identified (Guo et al. 2007). Several nanovirus-like DNA (DNA1) variants ranging in size from 1373–1385 nt were identified with a different isolates of MaYMV from Hainan, China (Guo and Xie 2008).

Malvastrum yellow vein Cambodia virus Taxonomic position Genus: Begomovirus

(MaYVCV)

Family: Geminiviridae

Geographical distribution MaYVCV infection in plants of Malvastrum coromandelianum was reported from Cambodia (Tang and He. unpublished; KP188831) Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2737 nt (KP188831 = NC_026763) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Malvastrum spp.

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Malvastrum yellow vein Honghe virus Taxonomic position Genus: Begomovirus

(MaYVHV)

Family: Geminiviridae

Geographical distribution MaYVHV infection in plants of Malvastrum coromandelianum was reported from China (FN552749). Symptoms and host(s) The virus-infected malvastrum plants exhibit yellow vein symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2740 nt (FN552749 = NC_030749) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Malvastrum yellow vein virus

(MaYVV)

M

Synonyms Kenaf leaf curl virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution MaYVV infection in plants of Malvastrum coromandelianum was reported from China (Zhou et al. 2003; Guo et al. 2008). Some isolates of MaYVV were previously known as Kenaf leaf curl virus and Malvastrum yellow vein Baoshan virus. Symptoms and host(s) The virus-infected malvastrum plants exhibit yellow vein symptoms. MaYVV has been isolated from Malvastrum coromandelianum, Hibiscus cannabinus, Sida acuta, Ageratum conyzoides, and Solanum lycopersicum. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2731 nt (AJ457824 = NC_004634) (Briddon 2001; Zhou et al. 2003; Brown et al. 2015; Zerbini et al. 2017).

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Malvastrum spp.

Malvastrum yellow vein Yunnan virus Taxonomic position Genus: Begomovirus

(MaYVYnV)

Family: Geminiviridae

Geographical distribution MaYVYnV infection in plants of Malvastrum coromandelianum was reported from Yunnan (China) (Jiang and Zhou 2005; Jiang et al. 2010). Symptoms and host(s) The virus-infected malvastrum plants exhibit yellow vein symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2747 nt (AJ786711 = NC_006631) (Briddon 2001; Jiang and Zhou 2005; Jiang et al. 2010; Brown et al. 2015; Zerbini et al. 2017).

Mesta yellow vein mosaic virus Taxonomic position Genus: Begomovirus

(MeYVMV)

Family: Geminiviridae

MeYVMV infection in plants of Malvastrum coromandelianum was reported from Pakistan (Anwar 2017). The virus-infected malvastrum plants exhibit yellow vein symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is not mechanically sap-transmissible. The virus is graft-transmissible. Kenaf leaf curl betasatellite is associated with the virus (Anwar 2017). For more details of MeYVMV, refer to Hibiscus sabdariffa.

Okra mosaic virus Taxonomic position Genus: Tymovirus

(OkMV)

Family: Tymoviridae

OkMV infection in plants of Malvastrum coromandelianum was reported from Nigeria (Atiri 1984). The virus is transmitted by beetle vectors in a semi-persistent manner, and is transmissible by mechanical sapinoculation, and by grafting. For more details of OkMV, refer to Abelmoschus esculentus.

Papaya leaf curl Guandong virus

(PaLCuGdV)

Synonyms Malvastrum leaf curl Guangdong virus (MaLCGdV)

Malvastrum spp.

Taxonomic position Genus: Begomovirus

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Family: Geminiviridae

MaLCGdV infection in plants of Malvastrum coromandelianum was reported from Guangdong Province of China (Wu et al. 2007). The virus-infected malvastrum plants exhibit leaf curl, vein swelling, and stunting symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Malvastrum leaf curl Guangdong betasatellite DNA molecule is associated with MaLCGdV, and consists of 1344 nt (KF912951) (Zhou 2013). For more details of PaLCuGdV, refer to Carica papaya.

Sida golden mosaic Florida virus Taxonomic position Genus: Begomovirus

(SiGMFlV)

Family: Geminiviridae

SiGMFlV infection in plants of Malvastrum coromandelianum was reported from Cuba (Fiallo-Olive et al. 2010). The virus-infected malvastrum plants exhibit golden mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of SiGMFlV, refer to Sida spp.

Sida yellow vein virus Taxonomic position Genus: Begomovirus

M

(SiYVV)

Family: Geminiviridae

SiYVV infection in plants of Malvastrum coromandelianum was reported from Cuba (Fiallo-Olive et al. 2012). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative nonpropagative manner. For more details of SiYVV, refer to Sida spp.

Tomato yellow leaf curl China virus

(TYLCCNV)

Synonyms Malvastrum yellow leaf curl virus (MaYLCV)

Taxonomic position Genus: Begomovirus

Family: Geminiviridae

MaYLCV infection in plants of Malvastrum coromandelianum was reported from China (Liu et al. 2009). The virus-infected malvastrum plants exhibit yellow leaf curl symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of TYLCCNV, refer to Solanum lycopersicum.

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Malvaviscus arboreus (Turk’s hat)

References Alabi OJ, Villegas C, Gregg L, Murray KD (2016) Complete nucleotide sequences of a new bipartite Begomovirus from Malvastrum spp. plants with bright yellow mosaic symptoms in South Texas. Arch Virol 161:1729–1733 Anwar S (2017) Association of Kenaf leaf curl betasatellite with Mesta yellow vein mosaic virus in naturally infected Malvastrum coromandelianum. Trop Plant Pathol 42(1):46–50 Atiri GI (1984) The occurrence and importance of Okra mosaic virus in Nigerian weeds. Ann Appl Biol 104:261–265 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer Index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Fiallo-Olive E, Martínez-Zubiaur Y, Moriones E, Navas-Castillo J (2010) Complete nucleotide sequence of Sida golden mosaic Florida virus and phylogenetic relationships with other begomoviruses infecting malvaceous weeds in the Caribbean. Arch Virol 155:1535–1537 Fiallo-Olive E, Navas-Castillo J, Moriones E, Martínez-Zubiaur Y (2012) Begomoviruses infecting weeds in Cuba: increased host range and a novel virus infecting Sida rhombifolia. Arch Virol 157:141–146 Graham AP, Stewart CS, Roye ME (2007) First report of a begomovirus infecting two common weeds: Malvastrum americanum and Sida spinosa in Jamaica. Plant Pathol 56:340 Graham AP, Martin DP, Roye ME (2010) Molecular characterization and phylogeny of two begomoviruses infecting Malvastrum americanum in Jamaica: evidence of the contribution of inter-species recombination to the evolution of malvaceous weed-associated begomoviruses from the Northern Caribbean. Virus Genes 40:256–266 Guo XJ, Xie Y (2008) Nanovirus-like DNA component associated with the Malvastrum yellow mosaic virus. Acta Virol 52:63–66 Guo X, Shi M, Zhou X (2007) Complete nucleotide sequences of Malvastrum yellow mosaic virus and its associated DNA beta molecule. Arch Virol 152(3):641–643 Guo W, Jiang T, Zhang X, Li GX, Zhou XP (2008) Molecular variation of satellite DNAb molecules associated with Malvastrum yellow vein virus and their role in pathogenicity. Appl Environ Microbiol 74:909–913 Huang JF, Zhou XP (2006) Molecular characterization of two distinct begomoviruses from Ageratum conyzoides and Malvastrum coromandelianum in China. J Phytopathol 154:648–653 Jiang T, Zhou X (2005) Molecular characterization of a distinct begomovirus species and its associated satellite DNA isolated from Malvastrum coromandelianum in China. Virus Genes 31(1):43–48 Jiang T, Liu P, Liao BL, Wu JX, Huang CJ (2010) Malvastrum yellow vein Yunnan virus is amonopartite begomovirus. Acta Virol 54(1):21–26 Liu P, Xie Y, Zhou XP (2009) Malvastrum coromandelianum is an alternative host of Tomato yellow leaf curl China virus. Plant Pathol 58:403 Wu J, Mugiira RB, Zhou X (2007) Malvastrum leaf curl Guangdong virus is a distinct monopartite begomovirus. Plant Pathol 56:771–776 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zhou X (2013) Advances in understanding begomovirus satellites. Annu Rev Phytopathol 51:357–381 Zhou X, Xie Y, Peng Y, Zhang Z (2003) Malvastrum yellow vein virus, a new Begomovirus species associated with satellite DNA molecule. Chin Sci Bull 48:2206–2210 Zia-Ur-Rehman M, Iqbal MJ, Hameed U, Haider MS (2018) First detection of Hollyhock leaf curl virus in Malvastrum coromandelianum in Pakistan. Plant Dis 102:256

Malvaviscus arboreus (Turk’s hat) Family: Malvaceae

Ornamental

Clerodendrum chlorotic spot dichorhavirus Taxonomic position Genus: Dichorhavirus

Family: Rhabdoviridae

(ClCSV)

Mandevilla spp.

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ClCSV infection in plants of Malvaviscus arboreus was reported from Brazil (Kitajima et al. 2003). The virus-infected Turk’s hat plants exhibit chlorotic spots on the foliage. The virus is transmitted by a mite vector Brevipalpus phoenicis, and also by mechanical sap-inoculation. For more details of ClCSV, refer to Clerodendrum spp.

Cotton leaf curl Multan virus Taxonomic position Genus: Begomovirus

(CLCuMuV)

Family: Geminiviridae

CLCuMuV infection in plants of Malvaviscus arboreus was reported from China (Tang et al. 2013). The virus-infected Turk’s hat plants exhibit upwards leaf curling, vein swelling, and vein dark green symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative nonpropagative manner. For more details of CLCuMuV, refer to Gossypium spp.

References Kitajima EW, Chagas CM, Rodrigues JCV (2003) Brevipalpus-transmitted plant virus and virus-like diseases: cytopathology and some recent cases. Exp Appl Acarol 30:135–160 Tang Y-F, He Z-F, Du Z-G, Han L-F, She X-M, Luo F-F (2013) Molecular characterization of the Cotton leaf curl Multan virus infecting Malvaviscus arboreus. Acta Phytopathol Sin 43(2):120–127

M Mandevilla spp. Family: Apocynaceae

Ornamental

Catharanthus mosaic virus Taxonomic position Genus: Potyvirus

(CatMV)

Family: Potyviridae

CatMV infection in plants of Mandevilla spp. was reported from the USA (Mollov et al. 2015). The virus-infected mandevilla plants showed mosaic symptoms, leaf deformation, premature leaf senescence, and vine dieback. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CatMV, refer to Catharanthus spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Mandevilla sanderi was reported from Italy (Davino et al. 2005, 2012). The virus-infected mandevilla plants exhibit symptoms of yellow rings in older leaves. The virus is

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Mandragora autumnalis (Autumn mandrake)

transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sapinoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

References Davino S, Bellardi MG, di Bella M, Davino M, Bertaccini A (2005) Characterization of a Cucumber mosaic virus isolate infecting Mandevilla sanderi (Hemsl.) Woodson. Phytopathol Mediterr 44:220–225 Davino S, Panno S, Rangel EA, Davino M, Bellardi MG, Rubio L (2012) Population genetics of Cucumber mosaic virus infecting medicinal, aromatic and ornamental plants from northern Italy. Arch Virol 157:739–745 Mollov D, Guaragna MA, Lockhart B, Rezende JAM, Jordan R (2015) First report of Catharanthus mosaic virus in Mandevilla in the United States. Plant Dis 99:165

Mandragora autumnalis (Autumn mandrake) Family: Solanaceae

Medicinal

Colombian datura virus Taxonomic position Genus: Potyvirus

(CDV)

Family: Potyviridae

CDV infection in plants of Mandragora autumnalis was reported from Italy (Pacifico et al. 2016). The virus-infected autumn mandrake plants exhibit flower and leaf mottle symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation, and by grafting. For more details of CDV, refer to Datura spp.

References Pacifico D, Crucitti D, Stigliano E, Ciuffo M, Vallino M, Carimi F (2016) First report of Colombian datura virus in Mandragora autumnalis in Sicily, Italy. Plant Dis 100:2338

Mangifera indica (Mango) Family: Anacardiaceae

Fruit crop

Mangifera indica latent virus

(MILV)

Taxonomic position MILV is a tentative member of the genus Benyvirus and family Benyviridae.

Manihot carthaginensis subspp. glaziovii (Tree cassava or Ceara rubber tree)

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Geographical distribution MILV infection in trees of Mangifera indica was reported from Israel (Sela et al. 2016). Symptoms and host(s) The virus-infected mango plants did not show any pathological symptoms on fruits or leaves. Virion properties and genome The genome has two molecules of positive-sense single-stranded RNA. RNA1 consists of 6725 nt (KU140662) and RNA2 of 2516 nt (KU140663). RNA1 encodes only one protein of 2,052 amino acids, and RNA2 encodes three different proteins: a putative helicase protein with a viral helicase-conserved domain that has a length of 352 amino acids, a putative coat protein with a Tobamovirus coat-conserved domain with a length of 182 amino acids, and a putative movement protein with a movement-protein-conserved viral domain and length of 134 amino acids (Gilmer et al. 2017).

References Gilmer D, Ratti C, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Benyviridae. J Gen Virol 98:1571–1572 Sela N, Luria N, Yaari M, Prusky D, Dombrovsky A (2016) Genome sequence of a potential new benyvirus isolated from mango RNA-seq data. Genome Announc 4(6):e01250–e01216. https://doi.org/10.1128/genomeA.01250-16

M Manihot carthaginensis subspp. glaziovii (Tree cassava or Ceara rubber tree) Family: Euphorbiaceae

Commercial crop

Cassava brown streak virus Taxonomic position Genus: Ipomovirus

(CBSV)

Family: Potyviridae

CBSV infection in trees of Manihot glaziovii was reported from Tanzania (Mbanzibwa et al. 2011). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CBSV, refer to Manihot esculenta.

References Mbanzibwa DR, Tian YP, Tugume AK, Mukasa SB, Tairo F, Kyamanywa S, Kullaya A, Valkonen JP (2011) Simultaneous virus-specific detection of the two cassava brown streak-associated viruses by RT-PCR reveals wide distribution in East Africa, mixed infections, and infections in Manihot glaziovii. J Virol Methods 171(2):394–400

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Manihot esculenta (Cassava)

Manihot esculenta (Cassava) Family: Euphorbiaceae

Tuber crop

African cassava mosaic Burkina Faso virus Taxonomic position Genus: Begomovirus

(ACMBFV)

Family: Geminiviridae

Geographical distribution ACMBFV infection in plants of Manihot esculenta was reported from Burkina Faso (Tiendrebeogo et al. 2012). Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, nonpropagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2770 nt (HE616777) (Brown et al. 2015; Tiendrebeogo et al. 2012; Zerbini et al. 2017).

African cassava mosaic virus

(ACMV)

Synonyms Cassava latent virus Taxonomic position Genus: Begomovirus

Family: Geminiviridae

Geographical distribution ACMV is found in plants of Manihot esculenta throughout tropical Africa, Zambia (Storey and Nichols 1938; Bock 1994a; Berrie et al. 1997; Thresh et al. 1998; Alabi et al. 2008a, 2011) and in adjacent islands, including Cape Verde, Sao Tome and Principe, Madagascar, Mauritius, Bioko, and Seychelles (Fauquet et al. 1988; Fargette et al. 1990; Thresh et al. 1994; Ahohuendo and Sarkar 1997; Calvert and Thresh 2002; Were et al. 2004; Anitha et al. 2011; Matic et al. 2012; Zinga et al. 2012; Chikoti et al. 2013; Harimalala et al. 2015; Mulenga et al. 2016) and also in, Mauritius (Zinga et al. 2012). Symptoms and host(s) The virus-infected cassava plants exhibit symptoms of leaf mosaic patterns that affect discrete areas and are determined at an early stage of leaf development (Patil and Fauquet 2009). Leaf chlorosis may be pale yellow or nearly white with only a shade of green, or just noticeable paler than normal. The chlorotic areas are usually clearly defined and vary in size from that of a whole leaflet to small flecks or spots. Leaflets may show a uniform mosaic pattern, or the mosaic pattern may be localized to a few areas, often at the leaf bases. Distortion, reduction in leaflet size, and general stunting can be secondary effects that are associated with symptom severity.

Manihot esculenta (Cassava)

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Natural infection of this virus was detected in leguminous plant species (Senna occidentalis, Leucaena leucocephala, and Glycine max), castor oil plant (Ricinus communis), a weed host (Combretum confertum), and a wild species of cassava (Manihot glaziovii) (Alabi et al. 2007, 2008b). Transmission The virus is transmitted by a whitefly vector, Bemisia tabaci, in a persistent manner. The virus is retained when the vector molts and not transmitted congenitally to the progeny of the vector (Dubern 1979, 1994; Fishpool and Burban 1994). The virus is transmissible by mechanical inoculation (with difficulty to cassava but more readily to Nicotiana benthamiana) and by grafting and not transmitted by seed (Bock and Guthrie 1978). The primary spread of this virus disease takes place through the use of cassava cuttings collected from infected plants. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2779 nt (J02057 = NC_001467) and DNA-B of 2724 nt (J02058 = NC_001468) (Stanley and Gay 1983; Bock and Woods 1983; Fauquet and Fargette 1990; Briddon et al. 1998; Fondong et al. 2000; Briddon 2001; Böttcher et al. 2004; Patil and Fauquet 2009; Scholthof et al. 2011; Brown et al. 2015; Zerbini et al. 2017). ACMV, which was referred as Cassava latent virus was the first geminivirus for which the first complete genome sequence was generated (Stanley and Gay 1983). The infectivity of the cloned DNAs of ACMV was subsequently demonstrated to prove the Koch's postulation of the etiological agent of the cassava mosaic disease (Briddon et al. 1998).

Cassava American latent virus Taxonomic position Genus: Nepovirus

(CsALV)

Family: Secoviridae

Geographical distribution CsALV was first described by Walter et al. (1989) from Manihot esculenta cultivars originating from Brazil and Guyana. The virus spreads in Brazil and Guyana. Symptoms and host(s) The virus-infected cassava plants may exhibit symptoms of mild mosaic. Generally no symptoms on virus-infected cassava are observed. Transmission No vector has been identified, and the virus is mechanically sap-transmissible, and host range is limited to species in the families Chenopodiaceae, Euphorbiaceae, and Solanaceae. Local lesions are produced on Chenopodium quinoa and Chenopodium amaranticolor. Virion properties and genome The virions are isometric, non-enveloped of two types, but similar in size 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3) (Lennon et al. 1987). The genome of the virus is composed of two RNAs (RNA-1 and RNA-2). RNA-2 was detected in the middle and the bottom nucleoprotein components and RNA-1 only in the bottom component (Walter et al. 1989; Mayo and

M

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Manihot esculenta (Cassava)

Robinson 1996; Sanfacon et al. 2009; Dunez and Le Gall 2011; Sanfacon 2015; Thompson et al. 2017). No sequence is currently available.

Cassava brown streak virus Taxonomic position Genus: Ipomovirus

(CBSV)

Family: Potyviridae

Geographical distribution CBSV infection in plants of Manihot esculenta was reported from Burundi, Guinea, Kenya, Malawi, Mozambique, Rwanda, Burundi, Congo, Tanzania, Uganda, Zanzibar, Zimbabwe, and Zambia (Storey 1936; Nichols 1950; Bock 1994b; Thresh et al. 1994; Legg and Raya 1998; Thresh and Mbwana 1998; Hillocks et al. 1999, 2002; Monger et al. 2001a, b; Calvert and Thresh 2002; Hillocks 2003; Hillocks and Jennings 2003; Alicai et al. 2007; Mware et al. 2009a; CABI/EPPO 2013a; Rwegasira et al. 2011; Mohammed et al. 2012; Patil et al. 2015). The virus is distributed widely in the highlands (>1000 m above the sea level) of the Lake Victoria zone in Uganda and Tanzania and also in the Indian Ocean coastal lowlands of Tanzania (Mbanzibwa et al. 2011a, b).

Symptoms and host(s) The virus-infected cassava plants develop leaf chlorosis, and roots become necrotic in storage. Leaf scars persist longer than usual after leaf drop, and brown lesions sometimes occur on young stems and as brown streaks on stems (Patil et al. 2015). The major symptoms observed were chlorotic leaf mottling, leaf chlorosis, vein-clearing, necrotic yellow spots, leaf collapse, reduced leaf sizes, interveinal chlorosis, and leaf curling. The characteristic yellow or necrotic vein-banding may enlarge and coalesce to form comparatively large, yellow patches. Tuberous root symptoms may also be present: these consist of dark-brown necrotic areas within the tuber and reduction in root size; lesions in roots can result in postharvest spoilage of the crop (Hillocks et al. 1999; Thresh and Hillocks 2003; Rwegasira and Rey 2012). The virus is also detected in the perennial species Manihot glaziovii in Tanzania, which revealed the first virus reservoir other than cassava (Mbanzibwa et al. 2011a). Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a non-persistent manner (Maruthi et al. 2005; Mware et al. 2009a, b). The virus is transmissible by mechanical sap-inoculation, and infects other Manihot spp. and some other hosts like Nicotiana spp. (Lister 1959). The virus is also transmissible by grafting. There is no evidence of transmission through true seeds.

Virion properties and genome The virions are non-enveloped, flexuous filaments and 800–950 nm long and 12–15 nm diameter. The genome consists of a single molecule of linear, positive-sense ssRNA of 9008 nt (GU563327 = NC_012698) (Monger et al. 2001a, 2010; Mbanzibwa 2011; Mbanzibwa et al. 2011a; Patil et al. 2015; Wylie et al. 2017).

Manihot esculenta (Cassava)

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Cassava common mosaic virus Taxonomic position Genus: Potexvirus

(CsCMV)

Family: Alphaflexiviridae

Geographical distribution CsCMV infection in plants of Manihot utilissima was first reported from Southern Brazil (Silberschmid 1938; Costa 1940). The virus spreads in Brazil, Colombia, Mexico, Peru, Taiwan, Ivory Coast, Sumatra, Indonesia, Cote d’Ivoire, and Tropical America from Mexico to Paraguay (Costa and Kitajima 1972; Aiton et al. 1988b; Lozano and Nolt 1989; Nolt et al. 1991; Thresh et al. 1994; Marys and Izaguirre-Mayoral 1995; Di Feo et al. 2015; Fernandez et al. 2017). Symptoms and host(s) The virus-infected cassava plants exhibit characteristic mosaic symptoms of yellowish areas on the leaf lamina, distortion, and stunting of the plant. On some of the virus-affected leaves, there are dark and light green patches that are delimited by veins. Symptoms are most severe during relatively cool periods. The virus is able to infect Manihot spp., Euphorbia prunifolia, Chenopodium amaranticolor, C. guinea, Malva parviflora, and Gossypium hirsutum. Transmission The virus is transmitted by means not involving a vector; and disseminated in infected cuttings. The virus is transmissible by mechanical inoculation to Nicotiana benthamiana (systemic host) and Chenopodium amaranticolor (local lesion host) and by grafting and not transmitted by true seed and pollen. Virion properties and genome The virions are filaments with a modal length of 495–525 nm and 15 nm wide. The genome consists of a single linear molecule of positive-sense single-stranded RNA of 6376 nt (U23414 = NC_001658) and comprises five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type and 18–27 kDa in size (Calvert et al. 1996; Adams et al. 2004).

Cassava frogskin-associated virus

(CsFSaV)

Taxonomic position CsFSaV is a tentative member of the genus Oryzavirus and family Reoviridae. Geographical distribution Cassava frogskin disease was first reported from Southern Colombia (Pineda et al. 1983). CsFSaVinfected plants of Manihot esculenta were reported from Amazonas region, Colombia, Brazil, Costa Rica, Argentina, Latin America, and Venezuela (Lozano and Nolt 1989; Chaparro-Martinez and Trujillo-Pinto 2001a; Calvert et al. 2008; Carvajal-Yepes et al. 2014; Di Feo et al. 2015). Symptoms and host(s) Most of the cassava genotypes infected by the virus do not show symptoms on the stems or leaves, but the roots are stunted and do not fill with starch. The lower stem may become enlarged. The root

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periderm and corky layer enlarge and form raised lip-shaped fissures. When the root symptoms are mild, the fissures often form a constricted ring around the root. A few cassava clones such as Secundina develop mosaic symptoms, and the plants are stunted. Transmission The virus is graft-transmissible. Virion properties and genome The virions are isometric particles of 70 nm which contain a genome consisting of ten dsRNA segments. Partial sequences of RNA3 (e.g. KF057036; 260 bp) and RNA4 (e.g. KF885741, 918 bp) are available.

Cassava green mottle virus Taxonomic position Genus: Nepovirus

(CsGMV)

Family: Secoviridae

Geographical distribution CsGMV infection in plants of Manihot esculenta spreads in the Pacific Islands, Australasia, and the Solomon Islands (Lennon and Aiton 1987; Lennon et al. 1986, 1987). Symptoms and host(s) The virus induces faint or distinct mottling and mosaic, symptoms in cassava. The affected leaves are often puckered with distorted margins. Plants often recover to give slightly stunted appearance but otherwise remain symptomless. Occasionally, plants are severely stunted. Transmission The possibility exists for transmission through nematodes, but none has been demonstrated yet. The virus is transmissible by mechanical sap-inoculation to cassava, Nicotiana clevelandii and Chenopodium quinoa. Virion properties and genome The virions are isometric, non-enveloped of two types, but similar in size 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is a bipartite, linear positive-sense, single-stranded RNA (Lennon et al. 1987; Sanfacon et al. 2009; Dunez and Le Gall 2011; Sanfacon 2015; Thompson et al. 2017). Only partial sequences of RNA 1 (MG581962; 930 nt) and RNA 2 (MG581963; 1749 nt) are available.

Cassava Ivorian bacilliform virus

(CIBV)

Taxonomic position CIBV is a tentative member of the genus Anulavirus and family Bromoviridae Geographical distribution CIBV infection in plants of Manihot esculenta was reported from Africa (Aiton et al. 1988a; Fargette et al. 1989; Fargette and Harrison 1990; Thresh et al. 1994).

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Symptoms and host(s) The virus-infected cassava plants do not exhibit any obvious symptoms. Transmission The virus is not transmitted by Myzus persicae. The virus is transmissible by mechanical sapinoculation, and members belonging to several (3–9) families are susceptible, which mostly show systemic chlorosis. Chenopodium murale is a local lesion host for this virus. Virion properties and genome The virions are bacilliform and non-enveloped, about 42–76  18 nm (Fargette et al. 1991). The genome consists of three segments of positive-sense ssRNA. RNA-1 consists of 3130 nt (KF742519 = NC_025482), RNA-2 of 2553 nt (KF742520 = NC_025483), and RNA-3 of 2564 nt (KF742521 = NC_025484) (Scott et al. 2014).

Cassava mosaic Madagascar virus Taxonomic position Genus: Begomovirus

(CMMGV)

Family: Geminiviridae

Geographical distribution CMMGV is a recombinant begomovirus characterized from Madagascar (Harimalala et al. 2012, 2015). Symptoms and host(s) The virus-infected cassava plants exhibit mosaic symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is a bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2769 nt (HE617299 = NC_017004) and DNA-B of 2741 nt (HE617300 = NC_017005) (Briddon 2001; Harimalala et al. 2012; Brown et al. 2015; Zerbini et al. 2017).

Cassava torrado-like virus

(CsTLV)

Taxonomic position CsTLV is a tentative member of the genus Torradovirus and family Secoviridae. Geographical distribution CsTLV infection in plants of Manihot esculenta was reported from Colombia and Argentina (CarvajalYepes et al. 2014).

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Manihot esculenta (Cassava)

Symptoms and host(s) Only recently it was found that complex virus infections are associated with root symptoms of Cassava frogskin disease (CFSD), that several novel viruses are part of that complex, and that neither CsFSaV nor a phytoplasma can induce CFSD symptoms or the associated leaf symptoms in single infections. CsTLV was associated with the leaf symptoms, but not the severe root symptoms of CFSD in both mixed and single infections (Carvajal-Yepes et al. 2014). Transmission The virus is transmitted by a whitefly vector in a persistent manner. The virus is graft-transmissible. Virion properties and genome The virus has a bipartite genome single-stranded positive sense RNA, and isometric virion particles of 25 nm; RNA 1 consists of 7256 nt (MF449522), and RNA 2 of 4489 nt (MF449523) (Verbeek et al. 2012; Carvajal-Yepes et al. 2014)

Cassava vein mosaic virus

(CsVMV)

Synonyms Cassava vein mottle virus Taxonomic position Genus: Cavemovirus

Family: Caulimoviridae

Geographical distribution CsVMV infection in plants of Manihot esculenta was first reported from Brazil in 1940 (Costa 1940). The virus has been reported from many states in Brazil (Lin and Kitajima 1980; Calvert et al. 1995; Calvert and Thresh 2002). Symptoms and host(s) The virus-infected cassava plants exhibit symptoms including chlorosis of the veins which can either appear as a chevron pattern or coalesce to form a ring-spot pattern. Some leaves show a mosaic pattern over the entire leaf. There is often leaf distortion, and sometimes the young leaves show epinasty. Expression of symptoms is variable, and symptoms may fade with age or not be expressed during certain times of the year. Transmission The virus is transmitted by vegetative propagation; no vector has been identified. Virion properties and genome The virions are isometric, non-enveloped, and 45–50 nm in diameter with no obvious surface structure (Kitajima and Costa 1980). The complete sequence of 8159 bp of an infectious clone of the doublestranded DNA genome of CsVMV was determined (U59751) and revealed a significant difference in genome organization when compared with a previous report (# U20341) (Calvert et al. 1995; de Kochko et al. 1998; Hohn 2011).

Manihot esculenta (Cassava)

Cassava virus C

1485

(CsVC)

Synonyms Cassava virus Q (CsVQ) Taxonomic position Genus: Ourmiavirus

Family: Botourmiaviridae

Geographical distribution CsVC was first isolated from plants of Manihot esculenta in Cote d’Ivoire and Malawi. The virus infection in cassava plants was reported from Cote d’Ivoire, Malawi, Zimbabwe, and Cameroon (Aiton et al. 1988a; Frison and Feliu 1991; Harrison et al. 1995; Thresh et al. 1994). Symptoms and host(s) The virus-infected cassava plants exhibit pronounced leaf fleck symptoms. Transmission The virus is mechanically sap-transmissible to Nicotiana benthamiana, Chenopodium amaranticolor, and C. quinoa. No vector is known. Virion properties and genome The virions are bullet-shaped, 18.5 nm in mean width, parallel-sided, and of several discrete lengths, 30 nm and 37 nm being the shortest and commonest lengths. The genome consists of three segments of positive-sense ssRNA. RNA-1 consists of 2740 nt (FJ157981 = NC_013111), RNA-2 of 1137 nt (FJ157982 = NC_013112), and RNA-3 of 958 nt (FJ157983 = NC_013113) (Rastgou et al. 2009; Milne and Accotto 2011; Turina et al. 2017).

Cassava virus X Taxonomic position Genus: Potexvirus

(CsVX)

Family: Alphaflexiviridae

Geographical distribution CsVX infection in plants of Manihot esculenta was reported from the north coast of Colombia and Venezuela (Lennon et al. 1986; Chaparro-Martinez and Trujillo-Pinto 2001b). Symptoms and host(s) The virus-infected cassava plants do not exhibit any symptoms (Chaparro-Martinez and Trujillo-Pinto 2001b). Transmission There is no known vector for this virus, but an aerial vector is suspected. The virus is transmitted through vegetative propagation. The virus is mechanically sap-transmissible to cassava (systemic infection), to Nicotiana benthamiana (systemic infection), or to Chenopodium quinoa and C. amaranticolor (local lesion hosts) (Lennon et al. 1986). The virus is not transmitted through seed.

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Virion properties and genome The virions are flexuous filaments with 495 nm in length and 13 nm in diameter. The genome consists of a single linear molecule of positive-sense ssRNA of 5879 nt (KY288509 = NC_034375) and encodes five ORFs; the replication-related proteins, the putative MPs (TGB), and the CP, respectively. Coat protein subunits are of one type and 18–27 kDa in size (Adams et al. 2004).

East African cassava mosaic Cameroon virus Taxonomic position Genus: Begomovirus

(EACMCV)

Family: Geminiviridae

Geographical distribution EACMCV infection in plants of Manihot esculenta was reported from Cameroon and Madagascar (Fondong et al. 1998; 2000; Alabi et al. 2008a; Fondong and Chen 2011; Harimalala et al. 2015). Symptoms and host(s) The virus-infected cassava plants exhibit mosaic symptoms. Cassava is the natural host of EACMCV. Natural infection of this virus was also found in legume species, Senna occidentalis and Leucaena leucocephala; a weed host, Combretum confertum; and a wild species of cassava, Manihot glaziovii (Alabi et al. 2007, 2008b). Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is a bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2802 nt (AF112354 = NC_004625) and DNA-B of 2741 nt (AF112355 = NC_004630) (Fondong et al. 2000; Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

East African cassava mosaic Kenya virus Taxonomic position Genus: Begomovirus

(EACMKV)

Family: Geminiviridae

Geographical distribution EACMKV infection in plants of Manihot esculenta was reported from Madagascar and Kenya (Bull et al. 2006; Harimalala et al. 2013, 2015). Symptoms and host(s) The virus-infected cassava plants exhibit mosaic symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner.

Manihot esculenta (Cassava)

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Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is a bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2797 nt (AJ717572 = NC_011583) and DNA-B of 2776 nt (AJ704971 = NC_011584) (Zhou et al. 1998; Briddon 2001; Bull et al. 2006; Brown et al. 2015; Zerbini et al. 2017).

East African cassava mosaic Malawi virus Taxonomic position Genus: Begomovirus

(EACMMV)

Family: Geminiviridae

Geographical distribution EACMMV infection in plants of Manihot esculenta was reported from Malawi (Africa) and Zambia (Zhou et al. 1998; Mulenga et al. 2015, 2016). Symptoms and host(s) The virus-infected cassava plants exhibit yellow mosaic and leaf distortion symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome consists of a single molecule of circular single-stranded DNA (DNA-A) of 2804 nt (AJ006460 = NC_022645) (Zhou et al. 1998; Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

East African cassava mosaic virus Taxonomic position Genus: Begomovirus

(EACMV)

Family: Geminiviridae

Geographical distribution EACMV infection in plants of Manihot esculenta was reported from the East African coast, Madagascar, Malawi, Mozambique, Zimbabwe, Western Kenya, Western Tanzania, Zambia, Nigeria, Togo, Guinea, Ivory Coast, Uganda, Angola, and Cameroon (Swanson and Harrison 1994; Gibson 1996; Ogbe et al. 1996, 1997, 1999, 2003; Zhou et al. 1997; Fondong et al. 1998; Winter 1998; Legg et al. 1999, 2001; Offei et al. 1999; Legg and Thresh 2000; Maruthi et al. 2002; Neuenschwamder et al. 2002; Bigirimana et al. 2004; Lava Kumar et al. 2009; Tiendrebeogo et al. 2009; Akinbade et al. 2010; Matic et al. 2012; Zinga et al. 2012; Harimalala et al. 2015; Mulenga et al. 2016). Symptoms and host(s) The virus-infected cassava plants exhibit mosaic, leaf distortion, and stunting symptoms.

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Manihot esculenta (Cassava)

Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The virus is mechanically sap-transmissible, and no virus transmission through true seed is reported. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is a bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2799 nt (AF126806 = NC_004674) and DNA-B of 2777 nt (AF126807 = NC_004676) (Briddon 2001; Hong et al. 1993; Bull et al. 2006; Brown et al. 2015; Zerbini et al. 2017).

East African cassava mosaic Zanzibar virus Taxonomic position Genus: Begomovirus

(EACMZV)

Family: Geminiviridae

Geographical distribution EACMZV infection in plants of Manihot esculenta was reported from Kenya and Zanzibar (Bull et al. 2003; Maruthi et al. 2004; Khan et al. 2013). Symptoms and host(s) The virus-infected cassava plants exhibit yellow mosaic and leaf distortion symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is a bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2785 nt (AF422174 = NC_004655) and DNA-B of 2763 nt (AF422175 = NC_004656) (Briddon 2001; Maruthi et al. 2004; Brown et al. 2015; Zerbini et al. 2017).

Indian cassava mosaic virus Taxonomic position Genus: Begomovirus

(ICMV)

Family: Geminiviridae

Geographical distribution ICMV infection in plants of Manihot esculenta was reported from India and Sri Lanka (Abraham 1956; Alagianagalingam and Ramakrishnan 1966; Malathi et al. 1987, 1988, 1989; Harrison et al. 1987; Aiton et al. 1988c; Harrison and Robinson 1988; Mathew 1989; Thresh et al. 1998; Saunders et al. 2002; Makeshkumar et al. 2005; Patil et al. 2005; Anitha et al. 2011). Symptoms and host(s) The virus-infected cassava plants exhibit mosaic and irregular chlorotic patches on leaves with distortion and stunting. Enations are sometimes observed on abaxial surfaces of leaves. Tuber splitting

Manihot esculenta (Cassava)

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is also observed. The disease significantly reduces stem girth and yield. Symptoms vary depending on the cultivar and season. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner (Antony et al. 2006, 2009). This virus is transmitted to species belonging to the Euphorbiaceae and Solanaceae through the whitefly vector (Mathew and Muniyappa 1993). The virus is transmissible with difficulty by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants, and nor through seed (Mathew and Muniyappa 1991). The primary spread of the disease is through infected planting materials, and secondary spread takes place through the whitefly vector (Duraisamy et al. 2013). Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is a bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2815 nt (Z24758 = NC_001932) and DNA-B of 2645 nt (Z24759 = NC_001933) (Hong et al. 1993; Briddon 2001; Saunders et al. 2002; Brown et al. 2015; Zerbini et al. 2017). The intergenic region of ICMV DNA contains three copies of a 41-nt sequence, called iteron sequences (Mathew and Muniyappa 1992).

South African cassava mosaic virus Taxonomic position Genus: Begomovirus

(SACMV)

Family: Geminiviridae

Geographical distribution SACMV infection in plants of Manihot esculenta was reported from South Africa, Swaziland (Rey and Thompson 1998; Berry and Rey 2001), and Madagascar (Ranomenjanahary et al. 2002). A strain of SACMV was detected in Zimbabwe, which indicated an expansion of its geographical range further north in mainland Africa, potentially through the trade of cassava between Southern Africa and Madagascar (Berrie et al. 1997; Briddon et al. 2004; Harimalala et al. 2015). Symptoms and host(s) The virus-infected cassava plants exhibit mosaic, leaf distortion, and stunting symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. The experimental hosts are Phaseolus vulgaris, Malva parviflora, and Nicotiana benthamiana which display symptoms of leaf curling, chlorosis, and stunting (Berrie et al. 2001). Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is a bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2800 nt (AF155806 = NC_003803) and DNA-B of 2760 nt (AF155807 = NC_003804) (Berrie et al. 1998, 2001; Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

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Soybean chlorotic blotch virus Taxonomic position Genus: Begomovirus

(SbCBV)

Family: Geminiviridae

SbCBV infection in plants of Manihot esculenta was reported from Cameroon and Togo (Leke et al. 2016). The virus-infected cassava plants exhibit mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of SbCBV, refer to Glycine max.

Sri Lankan cassava mosaic virus Taxonomic position Genus: Begomovirus

(SLCMV)

Family: Geminiviridae

Geographical distribution SLCMV infection in plants of Manihot esculenta was detected in India and Sri Lanka and Cambodia (Saunders et al. 2002; Dutt et al. 2005; Patil et al. 2005; Anitha et al. 2011; Wang et al. 2016). Symptoms and host(s) The virus-infected cassava plants exhibit symptoms of irregular yellow or green chlorotic foliar mosaic, leaf distortion, stunted growth, and decreased tuber yield (Rageshwari et al. 2013). Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner, and the virus is also mechanically sap-transmissible (Duraisamy et al. 2013). Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is a bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2755 nt (AJ314737 = NC_003861; AJ579307) and DNA-B of 2711 nt (AJ314738 = NC_003862; AJ579308) (Briddon 2001; Saunders et al. 2002; Patil et al. 2005; Dutt et al. 2005; Brown et al. 2015; Zerbini et al. 2017). SLCMV is capable of transreplicating betasatellites (Saunders et al. 2002; Patil and Fauquet 2010).

Ugandan cassava brown streak virus Taxonomic position Genus: Ipomovirus

(UCBSV)

Family: Potyviridae

Geographical distribution CBSV and UCBSV and their strains cause cassava brown streak disease in lowland and highland areas in Eastern Africa (CABI/EPPO 2013b; Patil et al. 2015). The viruses are known to infect Manihot esculenta in Tanzania and Uganda and most likely in all areas where the disease occurs. Isolates of UCBSV from the Lake Victoria zone were placed to two phylogenetic clusters in accordance with their origin in Uganda or Tanzania (Alicai et al. 2007; Mbanzibwa et al. 2011a, b; Mohammed et al. 2012).

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UCBSV has been detected in Burundi, a landlocked country in the African Great Lakes region of Southeast Africa (Bigirimana et al. 2011), and in Congo (Mulimbi et al. 2012). Symptoms and host(s) Symptoms of UCBSV are similar to those caused by CBSV in cassava. Challenge inoculation of CBSV and UCBSV isolates included necrotic local lesion, stunted growth, and mottling symptoms on N. rustica and N. tabacum which were not different between CBSV and UCBSV. However, the symptoms caused by the two viruses could be distinguished on N. benthamiana (Mbanzibwa et al. 2011a). In N. benthamiana, the UCBSV isolates consistently caused stunted growth, systemic mosaic, and rugosity in leaves but caused no local lesion symptoms (Winter et al. 2010), whereas CBSV isolates caused necrotic local lesion and severe systemic necrosis resulting to death of the plant. N. benthamiana therefore can be used as a differential host to identify these closely related ipomoviruses infecting cassava. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a non-persistent manner. The virus is also mechanically sap-transmissible and transmitted by grafting (Wagaba et al. 2013; Dombrovsky et al. 2014). Virion properties and genome The virions are flexuous filaments and 800–950 nm long and 11–20 nm diameter. The genome consists of a positive-sense ssRNA (Alicai et al. 2007). The genome of the highland isolate MLB3 is 9,069 nt (FJ039520) (Mbanzibwa et al. 2011a). The complete genome sequence of another isolate of UCBSV (Ug_23) collected from Namulonge, Uganda, in 2007 was determined, which contained 9070 nt (FJ185044 = NC_014791) (Monger et al. 2010; Winter et al. 2010; Abarshi et al. 2012).

West African Asystasia virus 1 Taxonomic position Genus: Begomovirus

(WAAV1)

Family: Geminiviridae

WAAV1 infection in plants of Manihot esculenta was reported from Cameroon and Togo (Leke et al. 2016). The virus-infected cassava plants exhibit mosaic symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative non-propagative manner. For more details of WAAV1, refer to Asystasia gangetica.

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Aiton MM, Lennon AM, Roberts IM, Harrison BD (1988a) Two new cassava viruses from Africa. In: Abstracts 5th international congress of plant pathology, Kyoto, p 43 Aiton MM, Roberts IM, Harrison BD (1988b) Cassava common mosaic potexvirus from mosaic-affected cassava in the Ivory Coast. Ann Rept 1987. Scottish Crop Research Institute, Dundee, p 191 Aiton MM, McGrath PF, Robinson DJ, Roberts IM, Harrison BD (1988c) Variation in Indian cassava mosaic geminivirus. In: Annual report 1987. Scottish Crop Research Institute, Dundee, p 191 Akinbade SA, Hanna R, Nguenkam A, Njukwe E, Fotso A, Doumtsop A, Ngeve J, Tenku STN, Kumar PL (2010) First report of the East African cassava mosaic virus-Uganda (EACMV-UG) infecting cassava (Manihot esculenta) in Cameroon. 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Marah macrocarpus (Wild cucumber)

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Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J Gen Virol 98:529–531 Thresh JM, Hillocks RJ (2003) Cassava mosaic and cassava brown streak diseases in Nampula and Zambesia provinces of Mozambique. Roots 8:10–15 Thresh JM, Mbwana MW (1998) Cassava mosaic and Cassava brown streak virus in Zanzibar. Roots 6:6–9 Thresh JM, Fargette D, Otim-Nape GW (1994) The viruses and virus diseases of cassava in Africa. Afr Crop Sci J 2:459–478 Thresh JM, Otim-Nape GW, Thankappan M, Muniyappa V (1998) The mosaic disease of cassava in Africa and India caused by whitefly-borne geminiviruses. Rev Plant Pathol 77:935–945 Tiendrebeogo F, Lefeuvre P, Hoareau M, Traore VSE, Barro N, Reynaud B, Traore AS, Konate G, Traore O, Lett J-M (2009) Occurrence of East African cassava mosaic virus-Uganda (EACMV-UG) in Burkina Faso. Plant Pathol 58:783 Tiendrebeogo F, Lefeuvre P, Hoareau M, Harimalala MA, Bruyn Ade, Villemot J, Traore VSE, Konate G, Traore AS, Barro N, Reynaud B, Traore O, Lett JM (2012) Evolution of African cassava mosaic virus by recombination between bipartite and monopartite begomoviruses. Virol J 9:67 Turina M, Hillman BI, Izadpanah K, Rastgou M, Rosa C, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Ourmiavirus. J Gen Virol 98:129–130 Verbeek M, Tang J, Ward LI (2012) Two generic PCR primer sets for the detection of members of the genus Torradovirus. J Virol Methods 185:184–188 Wagaba H, Beyene G, Trembley C, Alicai T, Fauquet CM, Taylor NJ (2013) Efficient transmission of Cassava brown streak disease viral pathogens by chip bud grafting. BMC Res Notes 6:516 Walter B, Ladeveze I, Etienne L, Fuchs M (1989) Some properties of a previously undescribed virus from cassava: Cassava American latent virus. Ann Appl Biol 115:279–289 Wang HL, Cui XY, Wang XW, Liu SS, Zhang ZH, Zhou XP (2016) First report of Sri Lankan cassava mosaic virus infecting cassava in Cambodia. Plant Dis 100:1029 Were HK, Winter S, Maiss E (2004) Occurrence and distribution of cassava begomoviruses in Kenya. Ann Appl Biol 145:175–184 Winter S (1998) Analysis of cassava samples from Guinea to study presence of viruses contributing to cassava mosaic disease (CMD). West Afr Seed Planting Mater News Lett 2:7–8 Winter S, Koerbler M, Stein B, Pietruszka A, Paape M, Butgereitt A (2010) Analysis of cassava brown streak viruses reveals the presence of distinct virus species causing cassava brown streak disease in East Africa. J Gen Virol 91:1365–1372 Wylie SJ, Adams M, Chalam C, Kreuze J, López-Moya JJ, Ohshima K, Praveen S, Rabenstein F, Stenger D, Wang A, Zerbini FM, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Potyviridae. J Gen Virol 98:352–354 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zhou X, Liu Y, Calvert L, Munoz C, Otim-Nape GW, Robinson DJ, Harrison BD (1997) Evidence that DNA-A of a geminivirus associated with severe cassava mosaic disease in Uganda has arisen by interspecific recombination. J Gen Virol 78:2101–2111 Zhou X, Robinson DJ, Harrison BD (1998) Types of variation in DNA-A among isolates of East African cassava mosaic virus from Kenya, Malawi and Tanzania. J Gen Virol 79:2835–2840 Zinga I, Harimalala M, De Bruyn A, Hoareau M, Mandakombo N, Semballa S, Reynaud B, Lefeuvre P, Lett JM (2012) East African cassava mosaic virus-Uganda (EACMV-UG) and African cassava mosaic virus (ACMV) reported for the first time in Central African Republic and Chad. Plant Dis 26:17

Marah macrocarpus (Wild cucumber) Family: Cucurbitaceae

Leafy vegetable

Wild cucumber mosaic virus Taxonomic position Genus: Tymovirus

(WCMV)

Family: Tymoviridae

M

1498

Maranta spp. (Prayer plants)

Geographical distribution WCMV was first reported in plants of Marah macrocarpus from California, USA, by Freitag (1952). The virus spreads in the USA and Australia (Allen and Fernald 1971). Symptoms and host(s) The virus-infected wild cucumber plants exhibit mild leaf chlorosis symptoms. Transmission The virus is transmitted by a beetle vector, Acalymma trivittata, in a semi-persistent manner. The virus is transmissible by mechanical sap-inoculation, and by grafting. The virus is not transmissible by contact between plants (Walters 1969). Virion properties and genome The virions are non-enveloped, isometric capsid with T = 3 icosahedral symmetry, about 30 nm in diameter. The genome consists of a single molecule of positive-sense ssRNA. The 30 -terminus has a tRNA-like structure. A partial genome sequence of 1379 nt is available (AF035633) (Martelli et al. 2002).

References Allen TC, Fernald EK (1971) Recovery and partial characterization of wild cucumber mosaic virus from Marah oreganus. Plant Dis Reptr 55:546–550 Freitag JH (1952) Seven virus diseases of cucurbits in California. Phytopathology 42:8 Martelli GP, Sabanadzovic S, Abou-Ghanem Sabanadzovic N, Edwards MC, Dreher T (2002) The family Tymoviridae. Arch Virol 147(9):1837–1846 Walters HJ (1969) Beetle transmission of plant viruses. Adv Virus Res 15:339–363

Maranta spp. (Prayer plants) Family: Marantaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

CMV infection in plants of Maranta spp. was reported by Hearon (1979). The virus-infected green prayer plants exhibit bright yellow mosaic patterns, ring-spot, and feathering on the leaves. Infected plants show symptoms for most of the production period, and on rare occasions growth reduction of some shoots occurs. The virus is transmitted by a number of aphids in a non-persistent manner, and also through mechanical sap-inoculation. The use of virus-infected stalk propagation is the primary mode of spread. For more details of CMV, refer to Cucumis sativus.

Marrubium vulgare (White horehound)

1499

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Maranta leuconeura was reported from Italy (Roggero et al. 1999). The virusinfected prayer plants exhibit chlorotic-necrotic spots and apical malformation symptoms. The virus is transmitted by the thrips vector, Frankliniella occidentalis in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

Sugarcane mosaic virus Taxonomic position Genus: Potyvirus

(SCMV)

Family: Potyviridae

SCMV was detected in the USA in bedding plants of Maranta leuconeura var. erythroneura (red-veined prayer plant) imported from Costa Rica (Baker et al. 2010). The virus-infected prayer plants showed mosaic on the upper leaf surface and chlorotic lesions on the underside of the leaf; plants lacked vigor, and most plants eventually died. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SCMV, refer to Saccharum officinarum.

M References Baker CA, Wilber LJ, Jones L (2010) A new host diagnosed with a strain of Sugarcane mosaic virus in Florida: red-veined prayer plant (Maranta leuconeura erythroneura). Plant Dis 94:378 Hearon SS (1979) A ringspot of prayer plant caused by a strain of Cucumber mosaic virus. Plant Dis Reptr 63:32–36 Roggero P, Ciuffo M, Dellavalle G, Gotta P, Gallo S, Peters D (1999) Additional ornamental species as hosts of Impatiens necrotic spot tospovirus in Italy. Plant Dis 83:967

Marrubium vulgare (White horehound) Family: Lamiaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Medicinal

(CMV)

Family: Bromoviridae

CMV infection in plants of Marrubium vulgare was reported from New Zealand (Fletcher 2001). The virus-infected white horehound plants do not exhibit obvious symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

1500

Masdevallia spp.

References Fletcher JD (2001) New hosts of Alfalfa mosaic virus, Cucumber mosaic virus, Potato virus Y, Soybean dwarf virus, and Tomato spotted wilt virus in New Zealand. N Z J Crop Hortic Sci 29:213–217

Masdevallia spp. Family: Orchidaceae

Ornamental

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Masdevallia spp. was reported from Germany and the United Kingdom (Lesemann and Koenig 1985; Hammond and Lawson 1988; Skelton et al. 2007). The virus-infected masdevallia plants may show a mosaic pattern of light and dark green areas, yellowing, lines of brown necrotic spots along the veins, ring-spot patterns of various kinds, and various necrotic spots often associated with pitting or sunken areas. Flowers may display color breaks and lines of necrotic spot following along the veins. Some plants show no symptoms at all (Hammond and Lawson 1988). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Odontoglossum ringspot virus Taxonomic position Genus: Tobamovirus

(ORSV)

Family: Virgaviridae

ORSV infection in plants of Masdevallia spp. was reported in the United Kingdom and Australia (Harju et al. 2011). The virus-infected masdevallia plants exhibit symptoms of necrotic spotting including some oval patterns. The virus is transmitted by means not involving a vector. The virus is transmissible by mechanical sap-inoculation. For more details of ORSV, refer to Odontoglossum grande.

References Hammond J, Lawson RH (1988) A strain of Bean yellow mosaic virus is aphid-transmitted from orchid. Acta Hortic 234:365–370 Harju VA, Skelton AL, Monger WA, Forde SMD, Daly M, Nixon T, Lawson R, Bennett S, Mumford RA (2011) A report of viruses, viroids and phytoplasmas found in ornamental plants from 1999–2007: findings from central science laboratory, UK. Acta Hortic 901:223–229 Lesemann DE, Koenig R (1985) Identification of Bean yellow mosaic virus in Masdevallia. Acta Hortic 164:347–354 Skelton A, Daly M, Nixon T, Harju V, Mumford RA (2007) First record of Bean yellow mosaic virus infecting a member of the orchid genus Dactylorhiza. Plant Pathol 56:344

Matthiola spp. (Stock)

1501

Matricaria chamomilla (Chamomile) Family: Asteraceae

Medicinal

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV infection in plants of Matricaria chamomilla was reported from Iran (Shahraeen et al. 2002). The virus-infected chamomile plants exhibit small necrotic spots, leaf yellowing, ring-spots, necrotic veinclearing, wilting, and dwarf symptoms. The virus is transmitted by thrips vectors in a persistentpropagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

References Shahraeen N, Ghotbi T, Mehraban AH (2002) Occurrence of Impatiens necrotic spot virus in ornamentals in Mahallet and Tehran provinces in Iran. Plant Dis 86:694

M Matthiola spp. (Stock) Family: Brassicaceae

Cauliflower mosaic virus Taxonomic position Genus: Caulimovirus

Ornamental

(CaMV)

Family: Caulimoviridae

CaMV infection in plants of Matthiola spp. was reported from Italy (Alioto et al. 1994). The virus-infected stock plants exhibit symptoms of severe stunting, mosaic, severe leaf deformation, flower break and greening, and floral abortion. The virus is transmitted by aphid vectors in a semi-persistent manner, and also by mechanical sap-inoculation. For more details of CaMV, refer to Brassica oleracea var. botrytis.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

1502

Maxillaria spp.

CMV infection in plants of Matthiola spp. was reported from Korea and northern Italy (JN593377) (Yoon et al. 1998; Davino et al. 2012). The virus-infected stock plants show prominent color break symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Turnip mosaic virus Taxonomic position Genus: Potyvirus

(TuMV)

Family: Potyviridae

TuMV infection in plants of Matthiola spp. was reported from the USA, Italy, Korea, and Iran (Tompkins 1934; Pound 1963; Bahar et al. 1985; Alioto et al. 1994; Farzadfar et al. 2005; Yoon et al. 1998). The virus-infected stock plants exhibit symptoms of prominent flower break, with foliar distortion and slight marbling. The virus is transmitted by the aphid vectors Brevicoryne brassicae and Myzus persicae in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TuMV, refer to Brassica rapa.

References Alioto D, Stavolone L, Aloj B (1994) Serious alterations induced by cauliflower mosaic virus (CaMV) and turnip mosaic virus (TuMV) on Matthiola incana. Inf Fitopatol 44:43–46 Bahar M, Danesh D, Dehghan M (1985) Turnip mosaic virus in stock plant. Iran J Plant Pathol 21:11 Davino S, Panno S, Rangel EA, Davino M, Bellardi MG, Rubio L (2012) Population genetics of Cucumber mosaic virus infecting medicinal, aromatic and ornamental plants from northern Italy. Arch Virol 157:739–745 Farzadfar SH, Ohshima K, Pourrahim R, Golnaraghi AR, Jalali S, Ahoonmanesh A (2005) Occurrence of Turnip mosaic virus on ornamental crops in Iran. Plant Pathol 54:261 Pound GS (1963) Resistance in Matthiola incana to the turnip mosaic virus. Phytopathology 53:1276–1279 Tompkins CM (1934) Breaking in stock (Mathiola incana), a virosis. Phytopathology 24:1137 Yoon JY, Choi HS, Ryu HY, Harm YI, Choi JK (1998) Color breaking syndrome of Matthiola incana caused by double infection of cucumber mosaic virus and turnip mosaic virus. Plant Pathol J 14(3):220–222

Maxillaria spp. Family: Orchidaceae

Ornamental

Orchid fleck dichorhavirus Taxonomic position Genus: Dichorhavirus

(OFV)

Family: Rhabdoviridae

OFV infection in plants of Maxillaria spp. was reported from Costa Rica (Freitas-Astua et al. 2002). The virus-infected maxillaria plants exhibit numerous yellow flecks and necrotic ring-spot lesions on

Mazus reptans (Creeping mazus)

1503

leaves. The virus is transmitted by Brevipalpus spp. mite vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of OFV, refer to Cymbidium spp.

References Freitas-Astua J, Moreira L, Rivera C, Rodriguez CM, Kitajima EW (2002) First report of Orchid fleck virus in Costa Rica. Plant Dis 86:1402

Mazus reptans (Creeping mazus) Family: Mazaceae

Ornamental

Alternanthera mosaic virus Taxonomic position Genus: Potexvirus

(AltMV)

Family: Alphaflexiviridae

AltMV infection in plants of Mazus reptans was reported from the USA (Henderson et al. 2014). The virus-infected creeping mazus plants exhibit mild mosaic symptoms. No vector is reported for this virus. The virus is readily sap-transmissible and easily spread by plant contact, vegetative propagation, and contaminated tools and hands. For more details of AltMV, refer to Alternanthera spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Mazus reptans was reported from the USA (Henderson et al. 2014). The virus-infected creeping mazus plants exhibit mild mosaic symptoms. The virus is transmitted by a number of aphid species in a non-persistent manner, and also by mechanical sap-inoculation, to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Ligustrum necrotic ringspot virus Taxonomic position Genus: Carlavirus

(LNRSV)

Family: Betaflexiviridae

LNRSV infection in plants of Mazus reptans was reported from the USA (Henderson et al. 2014). The virus-infected creeping mazus plants exhibit mild mosaic symptoms. The virus is presumed to be aphidtransmited, is mechanically sap-transmissible, and also by grafting. For more details of LNRSV, refer to Ligustrum spp.

M

1504

Medicago lupilina (Hop trefoil/Black medick)

References Henderson DC, Reinsel MD, Fischer KF, Hammond J (2014) First detection of Ligustrum necrotic ringspot virus, Cucumber mosaic virus, and Alternanthera mosaic virus in Mazus reptans in the United States. Plant Dis 98:1446

Medicago lupilina (Hop trefoil/Black medick) Family: Fabaceae

Forage crop

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

(AMV)

Family: Bromoviridae

AMV infection in plants of Medicago lupilina was reported from New Zealand (Fletcher 2001). The virus-infected hop trefoil plants do not exhibit obvious symptoms. The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Black medic leaf roll virus Taxonomic position Genus: Nanovirus

(BMLRV)

Family: Nanoviridae

Geographical distribution BMLRV infection in plants of Medicago lupilina was reported from Austria (Grigoras et al. 2014). Symptoms and host(s) The virus-infected hop trefoil plants exhibit symptoms of upward leaf rolling, sometimes with reddening and stunting. Transmission The virus is transmitted by aphid species Acyrthosiphum pisum and Aphis craccivora in a persistent but not in a propagative manner. The virus is retained when the vector molts, but does not multiply in the vector. The virus is not transmissible mechanically or through seeds. Virion properties and genome The virions are non-enveloped and spherical of about 17–20 nm in diameter with icosahedral symmetry. The genome consists of eight circular, positive-sense single-stranded DNA components, ranging in size from 1008 to 1019 nt and with a total genome size of 8016 nt (KC978949 – KC978956; NC_023299 – NC_023307). Each ssDNA segment has a common stem-loop region and each is encapsidated in a separate particle (Grigoras et al. 2014).

Medicago lupilina (Hop trefoil/Black medick)

Chickpea chlorotic stunt virus Taxonomic position Genus: Polerovirus

1505

(CpCSV)

Family: Luteoviridae

CpCSV infection in plants of Medicago lupilina was reported from Syria (Asaad et al. 2009). The virusinfected hop trefoil plants exhibit yellowing, reddening, and/or stunting symptoms. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of CpCSV, refer to Cicer arietinum.

Hop trefoil cryptic virus 1 Taxonomic position Genus: Unassigned

(HTCV-1)

Family: Partitiviridae

Geographical distribution HTCV-1 was first reported in plants of Medicago lupilina from Italy by Boccardo et al. (1987) and Luisoni et al. (1987). Symptoms and host(s) The virus-infected hop trefoil plants are symptomless. Transmission The virus is not transmissible by mechanical sap-inoculation, and no vector is involved. The virus is not transmissible by grafting. Transmission of this virus takes place vertically through infected seed (Boccardo et al. 1983). No sequence information is currently available. Virion properties and genome The virions are isometric, non-enveloped, and 30 nm in diameter. The genome typically consists of two dsRNA segments (Vainio et al. 2018).

Hop trefoil cryptic virus 2 Taxonomic position Genus: Betapartitivirus

(HTCV-2)

Family: Partitiviridae

Geographical distribution HTCV-2 was first reported in plants of Medicago lupilina from Italy by Boccardo et al. (1987) and Luisoni et al. (1987). Symptoms and host(s) The virus-infected hop trefoil plants are symptomless. Transmission The virus is not mechanically sap-transmissible and no insect vector is reported. The virus is not grafttransmissible. Transmission of this virus takes place through infected seed (Boccardo et al. 1983, 1987).

M

1506

Medicago lupilina (Hop trefoil/Black medick)

Virion properties and genome The virions are isometric, non-enveloped, and 30–35 nm in diameter. The genome consists of two doublestranded RNAs. RNA1 consists of 2431 bp (JX971980 = NC_021098) and RNA2 of 2349 bp (JX971981 = NC_021099) (Lesker et al. 2013; Vainio et al. 2018).

Hop trefoil cryptic virus 3 Taxonomic position Genus: Unassigned

(HTCV-3)

Family: Partitiviridae

Geographical distribution HTCV-3 was first reported in plants of Medicago lupilina from Italy by Boccardo et al. (1987) and Luisoni et al. (1987).

Symptoms and host(s) The virus-infected hop trefoil plants are symptomless.

Transmission The virus is not mechanically sap-transmissible and no insect vector is reported. The virus is not transmissible by grafting. Transmission of this virus takes place through infected seed (Boccardo et al. 1983, 1987).

Virion properties and genome The virions are isometric 30 nm in diameter. The genome typically consists of two dsRNA segments (Vainio et al. 2018). No sequence information is currently available.

References Asaad NY, Kumari SG, Haj-Kassem AA, Shalaby A-BA, Al-Shaabi S, Malhotra RS (2009) Detection and characterization of Chickpea Chlorotic Stunt Virus in Syria. J Phytopathol 157:756–761 Boccardo G, Lisa V, Milne RG (1983) Cryptic viruses in plants. In: Company RW, Bishop DHL (eds) Double-stranded RNA ‘viruses’. Elsevier, Amsterdam, pp 425–430 Boccardo G, Lisa V, Luisoni E, Milne RG (1987) Cryptic plant viruses. Adv Virus Res 32:171–214 Fletcher JD (2001) New hosts of Alfalfa mosaic virus, Cucumber mosaic virus, Potato virus Y, Soybean dwarf virus, and Tomato spotted wilt virus in New Zealand. N Z J Crop Hortic Sci 29:213–217 Grigoras I, Ginzo AIC, Martin DP, Varsani A, Romero J, Mammadov AC, Huseynova IM, Aliyev JA, Kheyr-Pour A, Huss H, Ziebell H, Timchenko T, Vetten H-J, Gronenborn B (2014) Genome diversity and evidence of recombination and reassortment in nanoviruses from Europe. J Gen Virol 95:1178–1191 Lesker T, Rabenstein F, Maiss E (2013) Molecular characterization of five betacryptoviruses infecting four clover species and dill. Arch Virol 158(9):1943–1952 Luisoni E, Milne RG, Accotto GP, Boccardo G (1987) Cryptic viruses in hop trefoil (Medicago lupulina) and their relationships to other cryptic viruses in legumes. Intervirology 28:144–156 Vainio EJ, Chiba S, Ghabrial SA, Maiss E, Roossinck M, Sabanadzovic S, Suzuki N, Xie J, Nibert M, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Partitiviridae. J Gen Virol 99:17–18

Medicago polymorpha (Burr clover)

1507

Medicago polymorpha (Burr clover) Family: Fabaceae

Weed host

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

(AMV)

Family: Bromoviridae

AMV infection in plants of Medicago polymorpha was reported from Australia (Jones and Nicholas 1992). The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sap-inoculation. The virus is found to be seed-transmitted in the range of 80–100% in Medicago polymorpha (Jones and Nicholas 1992; Pathipanowat et al. 1995). For more details of AMV, refer to Medicago sativa.

Bean leafroll virus Taxonomic position Genus: Luteovirus

(BLRV)

Family: Luteoviridae

BLRV infection in plants of Medicago polymorpha was reported from Australia (Schwinghamer et al. 1999). The virus-infected burr clover plants exhibit symptoms of yellowing or reddening, stunting, and stiffening. The virus is transmitted by several species of aphids, and the virus-vector relationship is of the circulative, non-propagative manner. The virus is not mechanically saptransmissible and is not transmitted through seed. For more details of BLRV, refer to Phaseolus vulgaris.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Medicago polymorpha was reported from Australia (Pathipanowat et al. 1995). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap inoculation. The virus is found to be seed-transmitted in the range of 0.3–1% in Medicago polymorpha (Pathipanowat et al. 1995). For more details of BYMV, refer to Phaseolus vulgaris.

Chickpea chlorotic stunt virus Taxonomic position Genus: Polerovirus

(CpCSV)

Family: Luteoviridae

M

1508

Medicago sativa (Alfalfa/Lucerne)

CpCSV infection in plants of Medicago polymorpha was reported from Syria (Asaad et al. 2009). The virus-infected burr clover plants exhibit yellowing, reddening, and/or stunting symptoms. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of CpCSV, refer to Cicer arietinum.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Medicago polymorpha was reported from Australia (Pathipanowat et al. 1995). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. The virus is found to be seed-transmitted up to 13% in Medicago polymorpha (Pathipanowat et al. 1995). For more details of CMV, refer to Cucumis sativus.

References Asaad NY, Kumari SG, Haj-Kassem AA, Shalaby A-BA, Al-Shaabi S, Malhotra RS (2009) Detection and characterization of Chickpea Chlorotic Stunt Virus in Syria. J Phytopathol 157:756–761 Jones RAC, Nicholas DA (1992) Studies on Alfalfa mosaic virus infection of burr medic (Medicago polymorpha) swards: seed-borne infection, persistence, spread and effects on productivity. Aust J Agric Res 43:697–715 Pathipanowat W, Jones RAC, Sivasithamparam K (1995) Studies on seed and pollen transmission of alfalfa mosaic, cucumber mosaic and bean yellow mosaic viruses in cultivars and accessions of annual Medicago species. Aust J Agric Res 46:153–165 Schwinghamer MW, Johnstone GR, Johnston-Lord CF (1999) First records of Bean leafroll luteovirus in Australia. Australas Plant Pathol 28:260

Medicago sativa (Alfalfa/Lucerne) Family: Fabaceae

Alfalfa cryptic virus 1 Taxonomic position Genus: Unassigned

Forage crop

(ACV-1)

Family: Partitiviridae

Geographical distribution ACV-1 infection in plants of Medicago sativa was reported from the Eurasian region, the Middle East, and Japan (Natsuaki et al. 1986; Accotto et al. 1990). Symptoms and host(s) The virus-infected alfalfa plants exhibit symptomless infections.

Medicago sativa (Alfalfa/Lucerne)

1509

Transmission There is no known vector for this virus. The virus is not transmissible by mechanical inoculation, not by grafting, and not by contact between plants; but is transmitted by seed and transmitted by pollen to the seed. Virion properties and genome The virions are isometric, non-enveloped, 30 nm in diameter, and rounded in profile, without a conspicuous capsomere arrangement. The genome typically consists of two dsRNA segments (Accotto et al. 1990; Vainio et al. 2018); no sequence information is currently available.

Alfalfa cryptic virus 2

(ACV-2)

Taxonomic position ACV-2 is a tentative member of the genus Betacryptovirus. Geographical distribution ACV-2 was first reported in plants of Medicago sativa from Japan by Boccardo et al. (1987). The virus spreads in Japan and Italy (Natsuaki et al. 1986). Symptoms and host(s) The virus-infected alfalfa plants exhibit symptomless infections. Transmission The virus is not transmissible by mechanical inoculation or by grafting but is transmitted by seed. Virion properties and genome The virions are isometric, 38 nm in diameter, and angular in profile. The genome is a double-strandedRNA; no sequence information is currently available.

Alfalfa dwarf cytorhabdovirus Taxonomic position Genus: Cytorhabdovirus

(ADV)

Family: Rhabdoviridae

Geographical distribution ADV infection in plants of Medicago sativa was reported from Argentina (Bejerman et al. 2011, 2015). Symptoms and host(s) The virus-infected alfalfa plants exhibit symptoms of shortened internodes, a bushy appearance, deformations, puckering, epinasty of leaflet blades, vein enations, and varying sized papillae on the adaxial leaflet surfaces. Transmission The virus is transmitted by the aphid vector, Aphis craccivora, in a persistent-propagative manner.

M

1510

Medicago sativa (Alfalfa/Lucerne)

Virion properties and genome The virions are enveloped, bacilliform, and measure 60–75 nm in diameter and 200–350 nm long. There are distinct surface projections (spikes) dispersed evenly over all the surface. The genome is a monopartite, negative-sense, single-stranded RNA of 14494 nt is unsegmented (KP205452 = NC_028237) and shows seven ORFs in antigenomic strand in the order of putative proteins N, P, P3, M, G, P6, and L (Dietzgen 2011; Bejerman et al. 2015; Walker et al. 2018).

Alfalfa enamovirus 1 Taxonomic position Genus: Enamovirus

(AEV-1)

Family: Luteoviridae

Geographical distribution AEV-1 infection in plants of Medicago sativa was reported from Argentina (Bejerman et al. 2016). A second isolate described as Alfalfa enamovirus 2 (AEV-2) was described from alfalfa in Sudan (Nemchinov et al. 2017). Symptoms and host(s) The virus-infected alfalfa plants exhibit dwarfing symptoms. Transmission The virus is transmitted by aphid vectors in a circulative, non-propagative manner, and not mechanically sap-transmissible. Virion properties and genome The virions are non-enveloped, spherical virion about 25 nm in diameter with T = 3 icosahedral symmetry composed of 180 CP proteins. The genome is a positive-sense, single-stranded RNA of 5726 nt (KU297983 = NC_029993) and shows 5 ORFs. The sequence of AEV-2 is of 5729 nt (KY985463).

Alfalfa leaf curl virus Taxonomic position Genus: Capulavirus

(ALCV)

Family: Geminiviridae

Geographical distribution ALCV infection in plants of Medicago sativa was reported from France (Roumagnac et al. 2015; Bernardo et al. 2016; Bejerman et al. 2018). Symptoms and host(s) The virus-infected alfalfa plants exhibit leaf curl symptoms. The virus naturally infects Medicago sativa (alfalfa) and can also experimentally infect Vicia faba (faba bean) and Nicotiana benthamiana (tobacco) causing a symptomless infection (Bernardo et al. 2016). Transmission The virus is transmitted by an aphid vector, Aphis craccivora (Roumagnac et al. 2015).

Medicago sativa (Alfalfa/Lucerne)

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Virion properties and genome The virions are twinned (geminate), incomplete icosahedra, and T = 1, 22  38 nm with a single coat protein. The reference genome is a monopartite circular, single-stranded DNA of 2745 nt (KP732474 = NC_027714), although sequence variants in the range of 2712–2769 nt are listed in GenBank (Varsani et al. 2014, 2017; Roumagnac et al. 2015; Zerbini et al. 2017; Bejerman et al. 2018).

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

(AMV)

Family: Bromoviridae

Geographical distribution AMV infection in plants of Medicago sativa was first reported from the USA (Weimer 1931). The virus occurs wherever alfalfa is grown (Ashby et al. 1979; Hampton and Weber 1983b; Forster et al. 1985; Bailiss and Ollennu 1986; Alhubaishi et al. 1987; Hiruki and Miczynski 1987; McLaughlin and Boykin 1988; Avgelis and Katis 1989; McLaughlin and Ensign 1989; Delgado Enquita and Luna Calvo 1992; Fidan 1992; Rahman and Peaden 1993; AL-Shahwan 2002; Kumari et al. 2005; van Leur and Kumari 2011; Massumi et al. 2012; Guy 2014; Al-Shahwan et al. 2017; Samarfard et al. 2017). Symptoms and host(s) The virus-infected alfalfa young leaves usually initially show small, round, green-yellow spots. Chlorotic spots enlarge in the form of stripes and rings or remain uneven. Spots on older leaves remain light yellow and whitish. The leaves of plants infected early are crinkled and show retarded growth. Decreased plant vigor results in shorter plants and plant death (Graham et al. 1972). The virus has a very wide host range infecting at least 700 species belonging to 71 families. Common hosts of the virus include alfalfa/lucerne (Medicago sativa), tomato (Solanum lycopersicum), potato (Solanum tuberosum), hot and sweet pepper (Capsicum spp.), beans (Phaseolus spp.), celery (Apium graveolens), and lettuce (Lactuca sativa) (Van Regenmortel and Pinck 1981). Transmission The virus is transmitted by 15 species of aphid vectors including, Myzus persicae, Acyrthosiphon pisum, and A. kondai, in a non-persistent manner (Garran and Gibbs 1982). The virus is mechanically sap-transmissible to a large number of host plants. The virus may be transmissible by grafting but is not transmissible by contact between plants. The virus is transmitted through seed at rates of up to 74% in alfalfa seeds from individual infected plants and up to 10% in commercial seed (Jones and Pathipanawat 1989; Bailiss and Offei 1990; Cali 1990; Jones 2004; Mando et al. 2004). The virus is transmitted by pollen to the seed. Virion properties and genome The virions are generally bacilliform, having a constant diameter of 19 nm, and varying from 30 to 56 nm in length, depending on the nucleic acid species encapsidated. The genome consists of three single-stranded RNA species and a fourth sub-genomic RNA from which the coat protein is translated. Transmission requires the presence of RNA-1 consisting of 3644 nt (L00163 = NC_001495); RNA-2, 2593 nt (X01572 = NC_002024); and RNA-3, 2037 nt (K02703 = NC_002025) together with the coat protein or RNA-4 (881 nt). Each genomic segment has a 30 tRNA-like structure and a 50 cap. The coat protein has a molecular mass of 24.25 kDa (Bol et al. 1971; Koper-Zwarthoff et al. 1979, 1980;

M

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Medicago sativa (Alfalfa/Lucerne)

Barker et al. 1983; Cornelissen et al. 1983a, b; Zuidema et al. 1983; Scott 2011; Bol and Linthorst 2011; Trucco et al. 2014).

Alfalfa virus F

(AVF)

Taxonomic position AVF is a tentative member of the genus Marafivirus, and family Tymoviridae Geographical distribution AVF was detected by high-throughput sequencing of Medicago sativa plants collected in southern France, with detection confirmed by RT-PCR (Nemchinov et al. 2018). Symptoms and host range No obvious symptoms differentiate an infected plant from healthy plants. Transmission No information on a vector is available; by analogy with other marafiviruses, a leafhopper or planthopper is suspected. Virion properties and genome The virions are spherical, of about 30 nm diameter, presumed to have T = 3 structure. The genome is a single molecule of positive-sense ssRNA, of 6818 nt (MG676465), most closely related to Medicago sativa marafivirus 1, another tentative member of the genus Marafivirus; however, the two sequences represent distinct species based on the criteria for demarcating marafivirus species (Nemchinov et al. 2018).

Alfalfa virus S Taxonomic position Genus: Allexivirus

(AVS)

Family: Alphaflexiviridae

Geographical distribution AVS infection in plants of Medicago sativa was reported from Sudan (Nemchinov et al. 2017b). Symptoms and host range The virus-infected plants exhibited chlorosis and stunting, but were also infected with peanut stunt virus and Alfalfa enamovirus1 (Nemchinov et al. 2017b). Transmission No information on a vector is available; by analogy with other allexiviruses, an eriophyoid mite is suspected. The virus is transmitted by mechanical sap inoculation. Use of infected vegetative propagative material is another source of virus spread. Virion properties and genome The virions are filamentous fexuous particles of 600–800 nm in length, 13–16 nm in diameter. The genome is a single molecule of positive sense ssRNA of 8349 nt (KY696659 = NC_034622), which

Medicago sativa (Alfalfa/Lucerne)

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encodes six open reading frames (respectively the viral replication protein, a triple gene block TGB1-3), a serine-rich 38.4 kDa protein, and a 29–31 kDa coat protein, but lack a 30 clear terminal ORF encoding a nucleic acid-binding protein typical of allexiviruses (Nemchinov et al. 2017b).

Bean common mosaic virus Taxonomic position Genus: Potyvirus

(BCMV)

Family: Potyviridae

BCMV infection in plants of Medicago sativa was reported from Central Anatolia and Saudi Arabia (Cali 1990; Al-Shahwan et al. 2017). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BCMV, refer to Phaseolus vulgaris.

Bean leafroll virus

(BLRV)

Synonyms Legume yellows virus; Pea leaf roll virus Taxonomic position Genus: Luteovirus

Family: Luteoviridae

BLRV infected alfalfa plants was reported from North America, Saudi Arabia, Argentina, Australia, and Iran (Cockbain and Gibbs 1973; Duffus 1979; Rahman and Peaden 1993; Kumari et al. 2005; van Leur and Kumari 2011; Massumi et al. 2012; Trucco et al. 2016; Al-Shahwan et al. 2017; Samarfard et al. 2017). The virus-infected alfalfa plants have a mild, transient yellowing of older leaves. The virus is confined to phloem and phloem parenchyma cells. The virus is transmitted by the pea aphid, Acyrthosiphon pisum, in a circulative, non-propagative manner. The virus is not mechanically saptransmissible. For more details of BLRV, refer to Phaseolus vulgaris.

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Medicago sativa was reported from the USA, Saudi Arabia, and Iran (McLaughlin and Boykin 1988; Bananej et al. 1995; Kumari et al. 2005; Massumi et al. 2012; Al-Shahwan et al. 2017). The virus-infected alfalfa plants exhibit symptoms of mosaic, leaf malformation and leaf mottling. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Beet western yellows virus Genus: Polerovirus

(BWYV)

Family: Luteoviridae

M

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Medicago sativa (Alfalfa/Lucerne)

BWYV infection in plants of Medicago sativa was reported from Yemen (Kumari et al. 2005). The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmitted by mechanical inoculation. For more details of BWYV, refer to Beta vulgaris.

Chickpea chlorotic stunt virus Taxonomic position Genus: Polerovirus

(CpCSV)

Family: Luteoviridae

CpCSV infection in plants of Medicago sativa was reported from Syria (Asaad et al. 2009). The virusinfected alfalfa plants exhibit symptoms of yellowing and reddening, and the plants are stunted. The virus is transmitted by aphid vectors in a circulative, non-propagative manner. The virus is not transmissible by mechanical sap-inoculation. For more details of CpCSV, refer to Cicer arietinum.

Clover yellow vein virus Taxonomic position Genus: Potyvirus

(ClYVV)

Family: Potyviridae

ClYVV infection in plants of Medicago sativa was reported from the Southern USA (McLaughlin and Boykin 1988). The virus is transmitted by several aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ClYVV, refer to Trifolium spp.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Medicago sativa was first reported from Yugoslavia, Saudi Arabia, and Iran (Babovic 1969; Singh and Singh 1983; Babovic and Matijevic 1985; Bananej et al. 1995; Al-Shahwan et al. 2017). The virus-infected alfalfa plants were widespread in distribution and exhibit symptoms of light-green mosaic/mottle on leaves. Leaflets are slightly malformed and plants have less vigor with retarded growth. The virus is transmitted by aphid vectors, Myzus persicae and Macrosiphon pisi, in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

Faba bean necrotic yellows virus Taxonomic position Genus: Nanovirus

(FBNYV)

Family: Nanoviridae

FBNYV was identified infecting plants of Medicago sativa in Yemen (Kumari et al. 2005). The virus is transmitted by aphid vectors in a persistent but not in a propagative manner. The virus is not transmitted by mechanical inoculation, not transmitted by contact between plants. For more details of FBNYV, refer to Vicia faba.

Medicago sativa (Alfalfa/Lucerne)

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Lucerne Australian latent virus Taxonomic position Genus: Nepovirus

(LALV)

Family: Secoviridae

Geographical distribution LALV was first reported in plants of Medicago sativa from Melbourne, Australia, by Blackstock (1978). The virus spreads in Australia and New Zealand (Taylor and Smith 1971; Ashby et al. 1979; Forster et al. 1985; Remah et al. 1986). Symptoms and host(s) The virus-infected lucerne (alfalfa) plants do not exhibit any symptoms. Transmission The virus is transmissible by mechanical sap-inoculation. The virus is transmitted by pollen to the seed. Seed-transmission up to 8% has been reported in lucerne (Taylor and Smith 1971). Virion properties and genome The virions are isometric and non-enveloped of two types but similar in size 25–30 nm in diameter and exhibit icosahedral symmetry (T = 1, pseudo T = 3). The genome is a bipartite, linear, positive-sense, single-stranded RNA, but no sequence information is available (Jones et al. 1979; Sanfacon et al. 2009; Dunez and Le Gall 2011; Sanfacon 2015; Thompson et al. 2017).

M Lucerne enation virus

(LEV)

Taxonomic position LEV is a tentative member of the family Rhabdoviridae. Geographical distribution LEV infection in plants of Medicago sativa was first reported from France (Alliot et al. 1972; Leclant et al. 1973). The virus spreads in France, Spain, Italy, Yugoslavia, Romania, Morocco, Saudi Arabia, and Eastern and Southern Europe (Cook and Wilton 1984; Marble 1984; Delgado Enquita and Luna Calvo 1992). Symptoms and host(s) The virus-affected lucerne (alfalfa) plants exhibit severe downward curling and puckering of leaflets and enations on the underside of the midveins of leaflets which become crinkled. Transmission The virus is transmitted by the cowpea aphid, Aphis craccivora, in a persistent manner. The virus is not transmissible either by sap or through seed. Virion properties and genome The virions are bullet-shaped and measure 82–89 nm in diameter and 250 nm in length. The genome is a linear, single-stranded RNA; no sequence information is available.

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Medicago sativa (Alfalfa/Lucerne)

Lucerne transient streak virus Taxonomic position Genus: Sobemovirus

(LTSV)

Family: Solemoviridae

Geographical distribution LTSV infection was first reported in plants of Medicago sativa from Australia. The virus spreads in Australia, Canada, New Zealand, and Saudi Arabia (Blackstock 1978; Ashby et al. 1979; Paliwal 1983; Forster et al. 1985; Rao and Hiruki 1985; Dall et al. 1990; Al-Shahwan et al. 2016a, 2017; Raza et al. 2017). Symptoms and host(s) The virus-infected lucerne (alfalfa) plants exhibit yellow streaks or flecks along the lateral veins of trifoliate leaves. The typical symptoms are transient streaks along the veinal areas (Rao and Hiruki 1985). The virus occurs naturally in alfalfa and in some other legume species including alsike clover, red clover, and sweet clover. Transmission No vectors have been reported for this virus. The virus is mechanically sap-transmissible. No seed and pollen transmission is recorded (Blackstock 1978; Forster and Jones 1979). Virion properties and genome The virions are isometric, non-enveloped, and 30 nm in diameter. The genome is a polycistronic, positive-sense, single-stranded RNA (ssRNA) of 4279 nt (JQ782213 = NC_001696). The genome consists of two overlapping ORFs, ORF2a, and ORF2b (Keese et al. 1983; Hull 1988; Paliwal 1984; Somera and Truve 2013; Somera et al. 2015).

Medicago sativa marafivirus 1

(MsMV-1)

Taxonomic position MsMV-1 is a tentative member of the genus Marafivirus, and family Tymoviridae. Geographical distribution The virus was identified by analysis in Korea of an alfalfa transcriptome from China (Kim et al. 2018). No validation by RT-PCR was therefore possible. Symptoms and host range Because of the nature of its detection, no symptom observations were possible. Transmission No information is available. Virion properties and genome The virions are presumed to be isometric, of c.30 nm diameter. The genome is a single molecule of positive-sense RNA of at least 6675 nt (MF443260) (Kim et al. 2018) which when compared to Alfalfa virus S appears to lack the extreme 50 sequence (Nemchinov et al. 2018).

Medicago sativa (Alfalfa/Lucerne)

Pea streak virus

1517

(PeSV)

Synonyms Alfalfa latent virus

Taxonomic position Genus: Carlavirus

Family: Betaflexiviridae

PeSV infection in plants of Medicago sativa was reported from the Czech Republic, Hungary, Saudi Arabia, and the USA (Veerisetty and Brakke 1977; Blackstock 1978; Hampton 1981; Hampton and Weber 1983a, b; Rahman and Peaden 1993; Nemchinov et al. 2015; Al-Shahwan et al. 2017). The virus-infected alfalfa plants exhibit brownish purple streaks on the stem, plant stunting, and necrosis symptoms. The complete sequence of the Alfalfa latent virus isolate of Nemchinov et al. (2015; KP784454) appears to lack a 30 -proximal nucleic acid binding protein, distinguishing it from other partial PeSV or another Alfalfa latent virus isolate. The virus is transmitted by the pea aphid, Acyrthosiphon pisum, in a non-persistent manner. This virus is mechanically sap-transmissible and is also transmitted through seed at a low rate (Hampton and Weber 1983a). For more details of PeSV, refer to Pisum sativum.

Peanut stunt virus Taxonomic position Genus: Cucumovirus

M

(PSV)

Family: Bromoviridae

PSV infection in plants of Medicago sativa was reported from Sudan, Spain, Iran, Saudi Arabia, and Southern United States (Diaz-Ruiz et al. 1979; Ahmed and El-Sadig 1985; McLaughlin and Boykin 1988; Bananej et al. 1998; Massumi et al. 2012; Al-Shahwan et al. 2017; Amid-Motlagh et al. 2017; Nemchinov et al. 2017b). The virus-infected alfalfa plants exhibit symptoms of bright leaf mottling to green-yellow mottle (Ahmed and Mills 1985). The virus is transmitted by the aphid vector, Aphis craccivora, in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts (Ahmed and El Sadig 1985). For more details of PSV, refer to Arachis hypogaea.

Red clover necrotic mosaic virus Taxonomic position Genus: Dianthovirus

(RCNMV)

Family: Tombusviridae

RCNMV infection in plants of Medicago sativa was reported from Victoria, Australia (Lynes et al. 1981). The virus-infected alfalfa plants exhibit necrosis and deformation symptoms. The virus is transmitted by a fungal vector Olpidium radicale, and also by mechanical sap-inoculation. For more details of RCNMV, refer to Trifolium spp.

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Medicago sativa (Alfalfa/Lucerne)

Red clover vein mosaic virus Taxonomic position Genus: Carlavirus

(RCVMV)

Family: Betaflexiviridae

RCVMV infection in plants of Medicago sativa was reported from Saudi Arabia, Canada, Europe, North America, and South America (Pesic and Hiruki 1987; McLaughlin and Boykin 1988; Rahman and Peaden 1993; Al-Shahwan et al. 2016b, 2017). The virus-infected alfalfa plants exhibit initial symptoms of yellow vein mosaic, and subsequently chlorosis occurs at the leaflet edges. Necrosis that occurs in interveinal areas results in premature plant death. There is a general reduction in the winter hardiness of diseased plants. The virus is transmitted by several aphid species, including the pea aphid, Acyrthosiphon pisum, and the green peach aphid, Myzus persicae, in a non-persistent manner. The virus is mechanically sap-transmissible; it is also transmitted through seed and pollen. For more details of RCVMV, refer to Trifolium spp.

Sweet clover necrotic mosaic virus Taxonomic position Genus: Dianthovirus

(SCNMV)

Family: Tombusviridae

SCNMV infection in plants of Medicago sativa was reported from Alberta, Canada (Inouye and Hiruki 1985; Hiruki 1986). The virus-infected alfalfa plants exhibit mosaic, ring-spots, and veinal necrosis symptoms. The virus is transmitted by the sweet clover weevil, Sitona cylindricollis (Hiruki 1986) and also by mechanical sap-inoculation. For more details of SCNMV, refer to Melilotus officinalis.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Medicago sativa was reported from eastern Washington, USA and Saudi Arabia (Kaiser et al. 1991; Al-Shahwan et al. 2017). The virus is transmitted by the thrips vectors, and they assist in the transfer of virus-infected pollen to the healthy plants. The virus is mechanically saptransmissible, but is not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

White clover mosaic virus Taxonomic position Genus: Potexvirus

(WClMV)

Family: Alphaflexiviridae

WClMV infection in plants of Medicago sativa was reported from the Southern USA and Saudi Arabia (McLaughlin and Boykin 1988; Cali 1990; Al-Shahwan et al. 2017). The virus-infected alfalfa plants exhibit mosaic symptoms. The virus is transmissible by mechanical inoculation but not by vectors. For more details of WClMV, refer to Trifolium spp.

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J Turk Phytopathol 21:15–20 Forster RLS, Jones AT (1979) Properties of Lucerne transient streak virus, and evidence of its affinity to Southern bean mosaic virus. Ann Appl Biol 93:181–189 Forster RLS, Morris-Krsinich BAM, Musgrave DR (1985) Incidence of Alfalfa mosaic virus, Lucerne Australian latent virus and Lucerne transient streak virus in lucerne crops in the North Island of New Zealand. N Z J Agric Res 28:279–282 Garran J, Gibbs A (1982) Studies on alfalfa mosaic virus and alfalfa aphids. Crop Pasture Sci 33:657–664 Graham JH, Kreitlow KW, Faulkner LR (1972) Diseases. Alfalfa science and technology, Agronomy Monograph No. 15. American Society of Agronomy, Madison, pp 497–523 Guy PL (2014) Viruses of New Zealand pasture grasses and legumes: a review. Crop Pasture Sci 65:841–853 Hampton RO (1981) Evidence suggesting identity between Alfalfa latent and Pea streak viruses. Phytopathology 71:223 Hampton RO, Weber KA (1983a) Pea streak virus transmission from alfalfa to peas: virus-aphid and virus-host relationships. Plant Dis 67:305–307 Hampton RO, Weber KA (1983b) Pea streak and alfalfa mosaic viruses in alfalfa: reservoir of viruses infectious to pisum peas. Plant Dis 67:308–310 Hiruki C (1986) Incidence and geographic distribution of Sweet clover necrotic mosaic virus in Alberta. Plant Dis 70:1129–1131 Hiruki C, Miczynski K (1987) Severe isolate of Alfalfa mosaic virus and its impact on alfalfa cultivars grown in Alberta. Plant Dis 71:1014–1018 Hull T (1988) The sobemovirus group. In: Koenig R (ed) The plant viruses. Polyhedral virions with monopartite RNA genomes, vol 3. Plenum Press, New York, pp 113–146 Inouye T, Hiruki C (1985) A new strain of Sweet clover necrotic mosaic virus isolated from alfalfa. Ann Phytopathol Soc Jpn 51:82 Jones RAC (2004) Occurrence of virus infection in seed stocks and three year – old pastures of lucerne (Medicago sativa). Aust J Agric Res 55:757–764 Jones RAC, Pathipanawat W (1989) Seed-borne Alfalfa mosaic virus infecting annual medics (Medicago spp.) in western Australia. Ann Appl Biol 115:263–277 Jones AT, Forster RLS, Mohamed NA (1979) Purification and properties of Australian lucerne latent virus, a seed-borne virus having affinities with nepoviruses. Ann Appl Biol 92:49–59 Kaiser WJ, Wyatt SD, Klein RE (1991) Epidemiology and seed transmission of two tobacco streak virus pathotype associated with seed increases of legume germ plasm in eastern Washington. Plant Dis 75:258–264 Keese P, Bruening G, Symons RH (1983) Comparative sequence and structure of circular RNAs from 2 isolates of Lucerne transient streak virus. FEBS Lett 159:185–190 Kim H, Park D, Hahn Y (2018) Identification of novel RNA viruses in alfalfa (Medicago sativa): an Alphapartitivirus, a Deltapartitivirus, and a Marafivirus. Gene 638:7–12 Koper-Zwarthoff EC, Brederode FT, Walstra P, Bol JF (1979) Nucleotide sequence of the 30 -noncoding region of Alfalfa mosaic virus RNA 4 and its homology with the genomic RNAs. Nucleic Acids Res 7(7):1887–1900 Koper-Zwarthoff EC, Brederode FT, Veeneman G, van Boom JH, Bol JF (1980) Nucleotide sequences at the 50 -termini of the Alfalfa mosaic virus RNAs and the intercistronic junction in RNA 3. Nucleic Acids Res 8(23):5635–5647

Medicago sativa (Alfalfa/Lucerne)

1521

Kumari SG, Muharram I, El-Pasha R, Al-Motwkel WA, Al-Ansi A (2005) First report of viruses that naturally infect alfalfa and fenugreek; and of Beet western yellows virus that infects legume crops in Yemen. Arab J Pl Prot 23:57 LeClant F, Alliot B, Signorent PA (1973) Transmission et epidemiologie de la maladie a enations de la Luzerne (Lev). Premiers resultats. Ann Phytopathol 5:441–445 van Leur JAG, Kumari SG (2011) A survey of lucerne in Northern New South Wales for viruses of importance to the winter legume industry. Australas Plant Pathol 40:180–186 Lynes EW, Teakle DS, Smith PR (1981) Red clover necrotic mosaic virus isolated from Trifolium repens and Medicago sativa in Victoria. Aust Plant Pathol 10:6–7 Mando JS, Kawas HZ, Makkouk KM, Kumari SG (2004) Forage legume viruses in Syria: economic importance and seed transmission. Arab J Pl Prot 22:122–127 Marble VL (1984) Report of consultancy on alfalfa and alfalfa seed production in Morocco, Saudi Arabia, the People’s Democratic Republic of Yemen, and Sudan, AGPC: MICS/85. Rome, FAO, p 82 Massumi H, Maddahian M, Heydarnejad J, Hosseini Pour A, Farahmand A (2012) Incidence of viruses infecting alfalfa in the Southeast and Central regions of Iran. J Agric Sci Technol 14:1141–1148 McLaughlin MR, Boykin DL (1988) Virus diseases of seven species of forage legumes in the Southeastern United States. Plant Dis 72:539–542 McLaughlin MR, Ensign RD (1989) Viruses detected in forage legumes in Idaho. Plant Dis 73:906–909 Natsuaki T, Natsuaki KT, Okuda S, Teranaka M, Milne RG, Boccardo G, Luisoni E (1986) Relationships between the cryptic and temperate viruses of alfalfa, beet and white clover. Intervirology 25:69–75 Nemchinov LG, Shao J, Postnikova OA (2015) Complete genome sequence of the Alfalfa latent virus. Genome Announc 3(2):e00250–e00215. https://doi.org/10.1128/genomeA.00250-15 Nemchinov LG, Grinstead SC, Mollov DM (2017a) First report and complete genome sequence of alfalfa enamovirus from Sudan. Genome Announc 5(27):e00531-17 Nemchinov LG, Grinstead SC, Mollov DM (2017b) Alfalfa virus S, a new species in the family Alphaflexiviridae. PLoS ONE 12(5):e0178222. Nemchinov LG, François S, Roumagnac P, Ogliastro M, Hammond RW, Mollov DS, Filloux D (2018) Characterization of alfalfa virus F, a new member of the genus Marafivirus. PLoS ONE 13(9):e0203477 Paliwal YC (1983) Identification and distribution in eastern Canada of Lucerne transient streak, a virus newly recognized in North America. Can J Plant Pathol 5:75–80 Paliwal YC (1984) Properties of a Canadian isolate of Lucerne transient streak virus and further evidence of similarity of the virus to sobemoviruses. Can J Plant Pathol 6:1–8 Pesic Z, Hiruki C (1987) Occurrence of Red clover vein mosaic virus (RCVMV) in alfalfa breeding lines. Phytopathology 77:1732 Rahman F, Peaden RN (1993) Incidence of viruses on Alfalfa in western North America. Plant Dis 77:160–162 Rao ALN, Hiruki C (1985) Occurrence and identification of Lucerne transient streak virus in Alberta, Canada. Plant Dis 69:610–612 Raza A, Al-Shahwan IM, Abdalla OA, Al-Saleh MA, Amer MA (2017) Lucerne transient streak virus; a recently detected virus infecting Alfafa (Medicago sativa) in Central Saudi Arabia. Plant Pathol J 33(1):43–52 Remah A, Jones AT, Mitchell MJ (1986) Purification and properties of lucerne Australian symptomless virus, a new virus infecting lucerne in Australia. Ann Appl Biol 109:307–315 Roumagnac P, Granier M, Bernardo P, Deshoux M, Ferdinand R, Galzi S, Fernandez E, Julian C, Abt I, Filloux D, Mesléard F, Varsani A, Blanc S, Martin DP, Peterschmitt M (2015) Alfalfa leaf curl virus: an aphid-transmitted geminivirus. J Virol 89:9683–9688 Samarfard S, Bejerman NE, Sharman M, Trucco V, Giolitti F, Dietzgen RG (2017) Development and validation of PCR assays for detection of alfalfa dwarf disease-associated viruses in Australian lucerne pastures. Australas Plant Pathol. https://doi.org/10.1007/s13313-017-0533-9 Sanfacon H (2015) In: Encyclopedia of Life Sciences (ed) Secoviridae: a family of plant picorna-like viruses with monopartite or bipartite genomes. Wiley, Chichester. https://doi.org/10.1002/9780470015902.a0000764.pub3 http:// www.els.net Sanfacon H, Wellink J, Le Gall O, Karasev A, van der Vlugt R, Wetzel T (2009) Secoviridae: a proposed family of plant viruses within the order Picornavirales that combines the families Sequiviridae and Comoviridae, the unassigned genera Cheravirus and Sadwavirus, and the proposed genus Torradovirus. Arch Virol 154(5):899–907 Scott SW (2011) Bromoviridae and allies. In: Encyclopedia of life sciences (ELS). Wiley, Chichester. https://doi.org/ 10.1002/9780470015902 Singh V, Singh RA (1983) Medicago sativa, a hitherto unrecorded natural reservoir of Cucumber mosaic virus. Indian J Mycol Plant Pathol 12:59–60 Somera M, Truve E (2013) The genome organization of lucerne transient streak and turnip rosette sobemoviruses revisited. Arch Virol 158:673–678 Somera M, Sarmiento C, Truve E (2015) Overview on Sobemoviruses and a proposal for the creation of the family Sobemoviridae. Viruses 7:3076–3115

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Megaskepasma erythrochlamys (Brazilian red-cloak)

Taylor RH, Smith PR (1971) Lucerne latent virus. Victorian Plant Research Institute Report No5. p 20 Thompson JR, Dasgupta I, Fuchs M, Iwanami T, Karasev AV, Petrzik K, Sanfaçon H, Tzanetakis IE, van der Vlugt R, Wetzel T, Yoshikawa N, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Secoviridae. J Gen Virol 98:529–531 Trucco V, de Breuil S, Bejerman N, Lenardon S, Giolitti F (2014) Complete nucleotide sequence of Alfalfa mosaic virus isolated from alfalfa (Medicago sativa L.) in Argentina. Virus Genes 48:562–565 Trucco V, de Breuil S, Bejerman N, Lenardon S, Giolitti F (2016) Bean leafroll virus (BLRV) in Argentina: molecular characterization and detection in alfalfa fields. Eur J Plant Pathol 146:207–212 Vainio EJ, Chiba S, Ghabrial SA, Maiss E, Roossinck M, Sabanadzovic S, Suzuki N, Xie J, Nibert M, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Partitiviridae. J Gen Virol 99:17–18 Van Regenmortel MHV, Pinck L (1981) Alfalfa mosaic virus. In: Kurstak E (ed) Handb Pl Virus Infec Comp Diag. Elsevier/North-Holland Biomedical Press, pp 415–421 Varsani A, Navas-Castillo J, Moriones E, Hernandez-Zepeda C, Idris A, Brown JK, Zerbini FM, Martin DP (2014) Establishment of three new genera in the family Geminiviridae: Becurtovirus, Eragrovirus and Turncurtovirus. Arch Virol 159:2193–2203 Varsani A, Roumagnac P, Fuchs M, Navas-Castillo J, Moriones E, Idris A, Briddon RW, Rivera-Bustamante R, Murilo Zerbini F, Martin DP (2017) Capulavirus and Grablovirus: two new genera in the family Geminiviridae. Arch Virol 162(6):1819–1831 Veerisetty V, Brakke MK (1977) Alfalfa latent virus a naturally occurring carlavirus in alfalfa. Phytopathology 67:1202–1206 Walker PJ, Blasdell KR, Calisher CH, Dietzgen RG, Kondo H, Kurath G, Longdon B, Stone DM, Tesh RB, Tordo N, Vasilakis N, Whitfield AE, ICTV Report Consortium (2018) ICTV virus taxonomy profile: Rhabdoviridae. J Gen Virol 99:447–448 Weimer JL (1931) Alfalfa mosaic. Phytopathology 21:122–123 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zuidema D, Bierhuizen MF, Cornelissen BJ, Bol JF, Jaspars EM (1983) Coat protein binding sites on RNA 1 of Alfalfa mosaic virus. Virology 125(2):361–369

Megaskepasma erythrochlamys (Brazilian red-cloak) Family: Acanthaceae

Ornamental

Broad bean wilt virus 1 Taxonomic position Genus: Fabavirus

(BBWV-1)

Family: Secoviridae

BBWV-1 infection in plants of Megaskepasma erythrochlamys was reported from Singapore (Koh et al. 2001). The virus-infected Brazilian red-cloak plants show mosaic and distortion symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BBWV-1, refer to Vicia faba.

Megakepasma mosaic virus Taxonomic position Genus: Unassigned

(MegMV)

Family: Closteroviridae

Melilotus albus (White sweet clover)

1523

Geographical distribution MegMV infection in plants of Megaskepasma erythrochlamys was reported by Mayo (2002a, b). Virion properties and genome The virions are flexuous filaments 12 nm wide and length ranging from 650 nm to over 2000 nm. The genome is a positive-sense, single-stranded RNA (Mayo 2002a, b). No sequence data is available.

References Koh LH, Cooper JI, Wong SM (2001) Complete sequences and phylogenetic analyses of a Singapore isolate of broad bean wilt fabavirus. Arch Virol 146:135–147 Mayo MA (2002a) Virus taxonomy – Houston 2002. Arch Virol 147(5):1071–1076 Mayo MA (2002b) A summary of taxonomic changes recently approved by ICTV. Arch Virol 147(8):1655–1656

Melampodium divaricatum (Butter daisy) Family: Asteraceae

Ornamental

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Melampodium divaricatum was reported from Louisiana, USA (Holcomb and Valverde 2000). The virus-infected butter daisy plants exhibit severe leaf mosaic, leaf malformation, and stunting symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Holcomb GE, Valverde RA (2000) First report of Oidium spp. Powdery Mildew and Tomato spotted wilt virus on Melampodium divaricatum. Plant Dis 84:1152

Melilotus albus (White sweet clover) Family: Fabaceae

Medicinal plant

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

M

1524

Melilotus albus (White sweet clover)

BYMV infection in plants of Melilotus albus was reported to be common in the USA, and elsewhere (Pierce and Wade 1933; Zaumeyer and Wade 1935; Schmidt 1977; Verhoyen and Meunier 1988). The virus-infected white sweet clover plants exhibit symptoms on leaves of small light yellow spots that later coalesce to form light-green zones, and then vein-clearing and leaf streaking occurs frequently. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Melilotus latent virus

(MeLV)

Synonyms Sweet clover latent virus Taxonomic position MeLV is a tentative member of the family Rhabdoviridae MeLV infection in plants of Melilotus albus was reported from the USA, Spain, and the Netherlands (Kitajima et al. 1969). The virus-infected white sweet clover plants do not exhibit any symptoms. No natural vector is reported for this virus. The virus is transmissible by grafting. For more details of MeLV, refer to Melilotus officinalis.

Tobacco streak virus Taxonomic position Genus: Ilarvirus

(TSV)

Family: Bromoviridae

TSV infection in plants of Melilotus albus was reported from Eastern Washington (Kaiser et al. 1982, 1991). The virus-infected white sweet clover plants do not exhibit symptoms. The virus is transmitted by thrips vectors, Thrips tabaci and Frankliniella occidentalis. The virus is seed-transmitted in M. albus plants to less than 3% (Kaiser et al. 1982). The virus is mechanically sap-transmissible, but not transmissible by contact between plants. For more details of TSV, refer to Nicotiana tabacum or Helianthus annuus.

References Kaiser WJ, Wyatt SD, Pesho GR (1982) Natural hosts and vectors of Tobacco streak virus in Eastern Washington. Phytopathology 72:1508–1512 Kaiser WJ, Wyatt SD, Klein RE (1991) Epidemiology and seed transmission of two tobacco streak virus pathotype associated with seed increases of legume germ plasm in eastern Washington. Plant Dis 75:258–264 Kitajima EW, Lauritis JA, Swift H (1969) Morphology and intracellular localization of a bacilliform latent virus in sweet clover. J Ultrastruct Res 29:141–150 Pierce WH, Wade BL (1933) Mosaic diseases affecting different legumes in relation to beans and peas. Phytopathology 23:562–564 Schmidt HE (1977) Leguminosen. In: Klinkowski M et al (eds) Pflanzliche Virologie, vol 3, 3rd edn. Akademie-Verlag, Berlin, pp 144–293 Verhoyen M, Meunier S (1988) Bean yellow mosaic virus (BYMV). In: Smith IM (ed) European Hand Pl Dis. Blackwell, London, pp 36–38 Zaumeter WJ, Wade BL (1935) The relationship of certain legume mosaics to bean. J Agric Res 51:715–749

Melilotus officinalis (Yellow sweet clover)

1525

Melilotus officinalis (Yellow sweet clover) Family: Fabaceae

Forage crop

Bean yellow mosaic virus Taxonomic position Genus: Potyvirus

(BYMV)

Family: Potyviridae

BYMV infection in plants of Melilotus officinalis was reported from the USA and England (Zaumeyer and Wade 1935; Verhoyen and Meunier 1988). The leaves of virus-infected yellow sweet clover plants exhibit small, light yellow spots that later coalesce to form light-green zones, and then vein-clearing and leaf streaking occurs frequently. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of BYMV, refer to Phaseolus vulgaris.

Melilotus latent virus

(MeLV)

Synonyms Sweet clover latent virus Taxonomic position MeLV is a tentative member of the family Rhabdoviridae. Geographical distribution MeLV infection in plants of Melilotus albus and M. officinalis was first reported from the USA, by Kitajima et al. (1969). The virus spreads in the USA, Spain, and the Netherlands (Knudson 1973). Symptoms and host(s) The virus-infected yellow sweet clover plants do not exhibit any symptoms. Transmission No natural vector is reported for this virus. The virus is transmissible by grafting. Virion properties and genome The virions are bullet-shaped and enveloped of 250–300 nm long and 80 nm wide. The genome is not segmented and consists of a single molecule of linear, single-stranded RNA. No sequence is available.

Sweet clover necrotic mosaic virus Taxonomic position Genus: Dianthovirus

(SCNMV)

Family: Tombusviridae

Geographical distribution SCMNV was first reported in plants of Melilotus officinalis from Athabasca, Alberta, Canada, by Hiruki et al. (1981). The virus spreads in Canada (Hiruki et al. 1984, 1989; Hiruki 1986).

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Melissa officinalis (Lemon balm)

Symptoms and host(s) The virus-infected yellow sweet clover plants exhibit systemic necrosis associated with leaf distortion and mosaic, resulting in severe stunting of the whole plant. Transmission The virus is transmitted by the sweet clover weevil vector, Sitona cylindricollis (Hiruki 1986). The virus is also transmissible by mechanical sap-inoculation (Hiruki et al. 1989). Virion properties and genome The virions are isometric, non-enveloped, 31–34 nm in diameter, and have T = 3 icosahedral symmetry. The genome is a bipartite, linear, single-stranded, positive-sense RNA. RNA1 consists of 3876 nt (L07884 = NC_003806) and RNA2 of 1449 nt (S46028 = NC_003807) (Hiruki 1987; Ge et al. 1992, 1993).

References Ge Z, Hiruki C, Roy KL (1992) A comparative study of the RNA-2 nucleotide sequences of two sweetclover necrotic mosaic virus strains. J Gen Virol 73:2483–2486 Ge Z, Hiruki C, Roy KL (1993) Nucleotide sequence of sweet clover necrotic mosaic dianthovirus RNA-1. Virus Res 28(2):113–124 Hiruki C (1986) Incidence and geographic distribution of Sweet clover necrotic mosaic virus in Alberta. Plant Dis 70:1129–1131 Hiruki C (1987) The dianthoviruses: a distinct group of isometric plant viruses with bipartite genome. Adv Virus Res 33:257–300 Hiruki C, Okuno T, Rao DV, Chen MH (1981) A new bipartite genome virus, sweet clover necrotic mosaic virus occurring in Alberta. A workshop paper on novel plant viruses. Abstracts, Fifth International Congress of Virology, p 235 Hiruki C, Rao DV, Chen MH, Okuno T, Figueiredo G (1984) Characterization of Sweet clover necrotic mosaic virus. Phytopathology 74:482–486 Hiruki C, Kudo K, Figueiredo G (1989) Transmission of Sweet clover necrotic mosaic virus. Proceedings of the Japan Academy. Ser B, Phys Biol Sci 65:243–247 Kitajima EW, Lauritis JA, Swift H (1969) Morphology and intracellular localization of a bacilliform latent virus in sweet clover. J Ultrastruct Res 29:141–150 Knudson DL (1973) Rhabdoviruses. J Gen Virol 20:105–130 Verhoyen M, Meunier S (1988) Bean yellow mosaic virus (BYMV). In: Smith IM (ed) European Hand Pl Dis. Blackwell, London, pp 36–38 Zaumeter WJ, Wade BL (1935) The relationship of certain legume mosaics to bean. J Agric Res 51:715–749

Melissa officinalis (Lemon balm) Family: Lamiaceae

Alfalfa mosaic virus Taxonomic position Genus: Alfamovirus

Medicinal

(AMV)

Family: Bromoviridae

AMV infection in plants of Melissa officinalis was reported from Northwest Bulgaria (Dikova and Rabenstein 2008). The virus-infected lemon balm plants exhibit symptoms of chlorotic spots on the leaves. The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Melochia spp.

1527

Tobacco rattle virus

(TRV)

Taxonomic position Genus: Tobravirus

Family: Virgaviridae

TRV infection in plants of Melissa officinalis was reported from Northwest Bulgaria (Dikova and Rabenstein 2008). The virus-infected lemon balm plants exhibit symptoms of necrotic spots on the stems. The virus is transmitted by nematode vectors and also by mechanical sap-inoculation. The virus is not transmissible by contact between plants. For more details of TRV, refer to Nicotiana tabacum.

Tulip virus X

(TVX)

Taxonomic position Genus: Potexvirus

Family: Alphaflexiviridae

TVX infection in plants of Melissa officinalis was reported from the USA (Tzanetakis et al. 2005). The virus-infected lemon balm plants exhibit bright variegation symptoms. The virus is transmitted by mite vectors and is also mechanical sap-transmissible. For more details of TVX, refer to Tulipa spp.

References Dikova B, Rabenstein F (2008) First report on the occurrence of Tobacco rattle virus and Alfalfa mosaic virus on Melissa officinalis L. in Bulgaria. Second congress of virology, proceedings and abstracts, Sofia, Bulgaria, 28–31 May, pp 341–346 Tzanetakis IE, Mackey IC, Martin RR (2005) Tulip virus X (TVX) associated with lemon balm (Melissa officinalis) variegation: first report of TVX in the USA. Plant Pathol 54:562

Melochia spp. Family: Malvaceae

Melochia mosaic virus Taxonomic position Genus: Begomovirus

Weed host

(MelMV)

Family: Geminiviridae

Geographical distribution MelMV infection in plants of Melochia spp. was reported from Brazil (Fiallo-Olivé et al. 2015). Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A consists

M

1528

Melothria spp. (Melothria liukiuensis, M. pendula)

of 2624 nt (KT201151 = NC_028140) and DNA-B of 2589 nt (KT201152 = NC_028141) (Briddon 2001; Brown et al. 2015; Fiallo-Olivé et al. 2015; Zerbini et al. 2017).

Melochia yellow mosaic virus Taxonomic position Genus: Begomovirus

(MelYMV)

Family: Geminiviridae

Geographical distribution MelYMV infection in plants of Melochia spp. was reported from Brazil (Fiallo-Olivé et al. 2015). Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci in a circulative non-propagative manner. Virion properties and genome Virions are twinned (geminate) incomplete icosahedra, T = 1, 22  38 nm with a single coat protein. The genome is bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2668 nt (KT201153 = NC_028142) and DNA-B of 2620 nt (KT201154 = NC_028143) (Briddon 2001; Brown et al. 2015; Fiallo-Olivé et al. 2015; Zerbini et al. 2017).

References Briddon RW (2001) Begomovirus. Geminiviridae. The Springer index of plant viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Fiallo-Olivé E, Zerbini FM, Navas-Castillo J (2015) Complete nucleotide sequences of two new begomoviruses infecting the wild malvaceous plant Melochia spp. in Brazil. Arch Virol 160:3161–3164 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Melothria spp. (Melothria liukiuensis, M. pendula) Family: Cucurbitaceae

Weed host

Squash vein yellowing virus Taxonomic position Genus: Ipomovirus

(SqVYV)

Family: Potyviridae

SqVYV infection in plants of Melothria pendula was reported from Florida (Adkins et al. 2011). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SqVYV, refer to Cucurbita pepo.

Mentha spp. (Mint)

1529

Watermelon mosaic virus Taxonomic position Genus: Potyvirus

(WMV)

Family: Potyviridae

WMV-1 infection in plants of Melothria pendula was reported from Florida (Adlerz 1972). The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of WMV, refer to Citrullus lanatus.

References Adkins S, Webster CG, Kousik CS, Webb SE, Roberts PD, Stansly PA, Turechek WW (2011) Ecology and management of whitefly-transmitted viruses of vegetable crops in Florida. Virus Res 159:110–114 Adlerz WC (1972) Melothria pendula plant infected with Watermelon mosaic virus – 1 as source of inoculum for cucurbits in collier county, Florida. J Econ Entomol 65:1303–1306

Mentha spp. (Mint) Family: Lamiaceae

Medicinal

Alfalfa mosaic virus

(AMV)

Taxonomic position Genus: Alfamovirus

Family: Bromoviridae

AMV infection in plants of Mentha spp. was reported from New Zealand, Germany, and the USA (Lovisolo and Luisoni 1963; Richter 1966; Gaborjanyi and Nagy 1972; Fletcher 2001; Tzanetakis et al. 2010). The virus-infected mint plants exhibit symptoms of interveinal chlorotic mottle of leaves and sometimes forming chlorotic spots and ring-spots. The virus is transmitted by several species of aphids in a non-persistent manner, and also by mechanical sap-inoculation. For more details of AMV, refer to Medicago sativa.

Arabis mosaic virus Taxonomic position Genus: Nepovirus

(ArMV)

Family: Secoviridae

ArMV infection in plants of Mentha spp. was reported from wherever the crop is grown (Tzanetakis et al. 2010). This virus is transmitted by nematode vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of ArMV, refer to Arabis hirsuta or Fragaria spp.

Chilli leaf curl India virus Taxonomic position Genus: Begomovirus

(ChiLCINV)

Family: Geminiviridae

M

1530

Mentha spp. (Mint)

ChiLCINV infection in plants of Mentha spicata was reported from India (Saeed et al. 2014, 2017). The virus-infected mint plants exhibit symptoms of yellow vein, leaf yellowing, mosaic, crinkling, and cupping. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, nonpropagative manner. For more details of ChiLCINV, refer to Capsicum annuum/Capsicum frutescens.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Mentha spp. was reported from Europe, China, Germany, Iran, and Turkey (Richter 1966; Vicchi and Bellardi 1988; Zhou et al. 1990; Sevik 2012; Forghani et al. 2014). The virusinfected mint plants exhibit mild to severe symptoms. The virus is transmitted by aphid vectors in a nonpersistent manner, and also by mechanical sap-inoculation. For more details of CMV, refer to Cucumis sativus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

INSV-infected plants of Mentha spp. were reported from Oregon (DeAngelis et al. 1994; Tzanetakis et al. 2010). INSV-infected mint plants exhibit symptoms of chlorosis, appearing first on terminals and moving progressively toward the base of the plant. Yellow leaf areas soon become necrotic, eventually resulting in the leaf death, followed by death of the entire aboveground portion of the plant. Occasional ring-spots are seen on the leaves. The virus is transmitted by the western flower thrips, Frankliniella occidentalis, in a persistent, propagative manner (DeAngelis et al. 1994) and also by mechanical sapinoculation to a wide range of host plants. For more details of INSV, refer to Impatiens spp.

Lychnis ringspot virus Taxonomic position Genus: Hordeivirus

(LRSV)

Family: Virgaviridae

LRSV infection in plants of Mentha spp. was reported from western Hungary (Beczner et al. 1992). The virus-infected mint plants exhibit mosaic symptoms. The virus is mechanically sap-transmissible. For more details of LRSV, refer to Silene spp. (Syn.) Melandrium spp. and Lychnis spp.

Mint vein banding-associated virus Taxonomic position Genus: Unassigned

(MVBaV)

Family: Closteroviridae

Mentha spp. (Mint)

1531

Geographical distribution MVBaV infection in plants of Mentha x gracilis cv. variegata was reported from the USA, and Canada (Tzanetakis et al. 2005a, 2010). Symptoms and host(s) The virus-infected mint plants exhibit yellow vein-banding symptoms. MVBaV was also detected in M. canadensis, M. spicata, and M. x piperita. Transmission The virus is transmitted by the mint aphid vector Ovatus crataegarius (Tzanetakis et al. 2005a). The virus is mechanically sap-transmissible and has a limited host range. Virion properties and genome The virions are flexuous filaments of about 1500 nm long and 12 nm wide. The genome consists of a single molecule of positive-sense, single-stranded RNA of 13,387 nt (KJ572575 = NC_038420) (Tzanetakis et al. 2005a).

Mint virus 1 Taxonomic position Genus: Closterovirus

(MV-1)

Family: Closteroviridae

Geographical distribution MV-1 infection in plants of Mentha x gracilis was reported from Oregon, USA (Tzanetakis et al. 2005b). Symptoms and host(s) The virus-infected mint plants exhibited leaf distortion and crinkling symptoms. MV-1 was also detected in M. spicata, and M. x piperita. Transmission The mint aphid, Ovatus crataegarius, transmitted this virus in a semi-persistent manner. The virus is graft-transmissible. Virion properties and genome The virions are filaments, non-enveloped, very flexuous, 1250–2200 nm long, and 10–13 nm in diameter. The genome is a monopartite, linear, single-stranded, positive-sense RNA of 15,450 nt (AY792620 = NC_006944) encoding nine proteins (Tzanetakis et al. 2005b; Agranovsky and Lesemann 2011).

Mint virus 2 Taxonomic position Genus: Vitivirus

(MV-2)

Family: Betaflexiviridae

M

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Mentha spp. (Mint)

Geographical distribution MV-2 infection in plants of Mentha x gracilis “Variegata” was reported from the USA (Tzanetakis et al. 2007). Symptoms and host(s) The virus-infected mint plants did not exhibit typical “Variegata” symptoms. Transmission The virus is transmitted by the mint aphid, Ovatus crataegarius, in the presence of a helper virus (presumed to be MV-1). The virus is not mechanically sap-transmissible (Tzanetakis et al. 2007). Virion properties and genome The virions are helically constructed flexuous filaments 725–825  12 nm, showing distinct crossbanding. The genome is a single molecule of positive-sense ssRNA of c. 7.6 kb; a partial sequence of 3897 nt is available (AY913795) and consists of five slightly overlapping ORFs which encode, in order, the replication-related proteins, a c.24 K protein with unknown functions, a MP of the “30 K” superfamily type, the CP, and a small protein (10–14 K) with nucleotide-binding properties. Coat protein subunits are of one type and 18–21.5 kDa in size (Adams et al. 2004).

Mint virus X Taxonomic position Genus: Potexvirus

(MVX)

Family: Alphaflexiviridae

Geographical distribution MVX infection in variegated plants of Mentha spp. was reported from the USA (Tzanetakis et al. 2006). Symptoms and host(s) The virus-infected mint plants do not exhibit any symptoms. Transmission The virus is not readily transmitted by mechanical sap-inoculation. No vector is known for this virus, but transmission may require presence of a helper virus. Virion properties and genome The virions are flexuous filaments 475–580 nm in length and 13 nm in diameter. The genome consists of a single molecule of a linear, positive-sense, single-stranded RNA of 5914 nt (AY789138 = NC_006948) and comprise five ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), and the CP. Coat protein subunits are of one type and 18–27 kDa in size (Adams et al. 2004; Tzanetakis et al. 2006).

Pedilanthus leaf curl virus

(PeLCV)

Synonyms Tomato leaf curl Pakistan virus (ToLCPKV)

Mentha spp. (Mint)

Taxonomic position Genus: Begomovirus

1533

Family: Geminiviridae

ToLCPKV/PeLCV infection in plants of Mentha spicata cv. viridis was reported from India (Samad et al. 2009). The virus-infected mint plants exhibit symptoms of mosaic, yellowing, leaf curling, crinkling, and retarded growth. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. For more details of PeLCV, refer to Euphorbia tithymaloides (Syn.) Pedilanthus tithymaloides.

Strawberry latent ringspot virus Taxonomic position Genus: Unassigned

(SLRSV)

Family: Secoviridae

SLRSV infection in plants of Mentha x gracilis cv. variegata was reported from the USA, and previously from wild Mentha species (Postman et al. 2004; Tzanetakis et al. 2010). The virus-infected mint plants do not exhibit any symptoms. The virus is transmitted by the nematode vectors, Xiphinema diversicaudatum and X. coxi and also by mechanical sap-inoculation. For more details of SLRSV, refer to Fragaria spp.

M Tobacco mosaic virus Taxonomic position Genus: Tobamovirus

(TMV)

Family: Virgaviridae

TMV infection in plants of Mentha x gracilis was reported from India (Samad et al. 1994, 2000). The virus-infected mint plants exhibit mosaic, green vein-banding, deformation of leaves, and poor stunted growth. There is no known vector for this virus. The virus is mechanically saptransmissible and also through contact between plants. For more details of TMV, refer to Nicotiana tabacum.

Tobacco ringspot virus Taxonomic position Genus: Nepovirus

(TRSV)

Family: Secoviridae

The virus infection in plants of Mentha  verticillata was reported from the USA (Stone et al. 1962; Tzanetakis et al. 2010). The virus-infected mint plants exhibit symptoms of mild vein-banding, stunting, and rosetting. The virus is transmitted by the nematode vector, Xiphinema spp., in a nonpersistent manner, and also by mechanical sap-inoculation. For more details of TRSV, refer to Nicotiana tabacum.

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Mentha spp. (Mint)

Tomato aspermy virus Taxonomic position Genus: Cucumovirus

(TAV)

Family: Bromoviridae

TAV infection in plants of Mentha spicata was reported from China (Zhou et al. 1990). The virusinfected mint plants exhibit symptoms of mosaic and distorted leaves. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of TAV, refer to Solanum lycopersicum.

Tomato leaf curl Karnataka virus Taxonomic position Genus: Begomovirus

(ToLCKaV)

Family: Geminiviridae

ToLCKaV infection in plants of Mentha arvensis, M. piperita, and M. spicata was reported from India (Borah et al. 2010). The virus-infected mint plants exhibit leaf deformity symptoms. The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner, and was also mechanically transmissible to Nicotiana benthamiana. For more details of ToLCKaV, refer to Solanum lycopersicum.

Tomato leaf curl Patna virus Taxonomic position Genus: Begomovirus

(ToLCPatV)

Family: Geminiviridae

ToLCPatV infection in plants of Mentha piperita was reported from India (Saeed and Samad 2016). The virus-infected mint plants exhibit symptoms of yellow mosaic, curling of leaves, and stunted growth. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. For more details of ToLCPatV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Mentha spp. was reported from Bulgaria, Italy, and Oregon, USA (Sether et al. 1991; Allen 1992; Grieco et al. 2000; Dikova 2011); however, some of these reports may refer to Impatiens necrotic spot virus, which was initially described as TSWV-Impatiens strain. The virusinfected mint plants exhibit symptoms of stunting and general decline. Newly mature, dark green leaves developed bright yellow, irregular mottling. Older leaves became bronze and then, often, completely yellow and developed irregular, brownish-gray, sunken lesions. The virus is transmitted by thrips vectors, such as Thrips tabaci, Frankliniella schultzei, F. fusca, F. occidentalis, and Scirtothrips dorsalis, in a persistent-propagative manner, and also by mechanical sap-inoculation to a wide range

Mentha spp. (Mint)

1535

of host plants. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060 Agranovsky AA, Lesemann D-E (2011) Closterovirus. Closteroviridae. In: The Springer Index of Viruses. Springer, New York, pp 327–333. https://doi.org/10.1007/978-0-387-95919-1_50 Allen TC. (1992) Tomato spotted wilt virus in mint. Mint Industry Research Council (MIRC) Proceedings Beczner L, Hamilton RI, Rochon DM (1992) Properties of the Mentha strain of Lychnis ringspot virus. Intervirology 33:49–56 Borah BK, Cheema GS, Gill CK, Dasgupta I (2010) A geminivirus-satellite complex is associated with leaf deformity of Mentha (mint) plants in Punjab. Indian J Virol 21:103–109 DeAngelis JD, Sether DM, Rossignol PA (1994) Transmission of impatiens necrotic spot virus in peppermint by western flower thrips (Thysanoptera: Thripidae). J Econ Entomol 87:197–201 Dikova B (2011) Tomato spotted wilt virus on some medicinal and essential oil-bearing plants in Bulgaria. Bulg J Agric Sci 17:306–313 Fletcher JD (2001) New hosts of Alfalfa mosaic virus, Cucumber mosaic virus, Potato virus Y, Soybean dwarf virus, and Tomato spotted wilt virus in New Zealand. N Z J Crop Hortic Sci 29:213–217 Forghani D, Mosahebi GM, Habibi-Koohi MM (2014) Characterization of Cucumber mosaic virus from mint in Tehran province. Int J Adv Biol Biomed Res 24:985–992 Gaborjanyi R, Nagy F (1972) Virus and mycoplasma diseases of cultivated medicinal plants in Hungary. Herba-Hung 11:39–51 Grieco PD, Conte D, Munno I, Nuzzaci M, de Stradis A (2000) Tomato spotted wilt virus (TSWV) on weeds and wild plants in Metapontino, Basilicata. Informatore Fitopatologico 50:43–46 Lovisolo O, Luisoni E (1963) A new virosis of peppermint and the presence in this plant of a virus inhibitor. Atti Acad Torino 98:213–225 Postman JD, Tzanetakis IE, Martin RR (2004) First report of Strawberry latent ringspot virus in Mentha sp. from North America. Plant Dis 88:907 Richter J (1966) Virus diseases in peppermint (Mentha piperita L.) in the German Democratic Republic. Pharmazie 21:373–377 Saeed ST, Samad A (2016) First detection of a monopartite Tomato leaf curl Patna virus infecting Mentha piperita in India. Plant Dis 100:2340 Saeed ST, Khan A, Kumar B, Ajayakumar PV, Samad A (2014) First report of Chilli leaf curl India virus infecting Mentha spicata (Neera) in India. Plant Dis 98:164 Saeed ST, Kumar B, Shasany AK, Samad A (2017) Molecular identification of Chilli leaf curl India virusalong with betasatellite molecule causing leaf curl disease of menthol mint (Mentha arvensis var. Kosi) in India. J Gen Plant Pathol 83:333–336 Samad A, Zaim M, Ajayakumar PV (1994) An outbreak of mosaic disease on mint (Mentha cardiaca Baker) in India. Indian J Plant Pathol 12:1–4 Samad A, Zaim M, Ajayakumar PV, Garg ID (2000) Isolation and characterization of a TMV isolate infecting scotch spearmint (Mentha gracilis Sole) in India. J Plant Dis Protect 107:649–657 Samad A, Gupta MK, Shasany AK, Ajayakumar PV, Alam M (2009) Begomovirus related to Tomato leaf curl Pakistan virus newly reported in Mentha crops in India. Plant Pathol 58:404 Sether DM, DeAngelis JD, Rossignol PA (1991) First report of Tomato spotted wilt virus in peppermint (Mentha piperita). Plant Dis 75(6):644 Sevik MA (2012) Natural occurrence of Cucumber mosaic virus infecting water mint (Mentha aquatica) in Antalya and Konya, Turkey. Acta Bot Croat 71(1):187–193 Stone WJ, Mins GI, Bergeson GB (1962) A new disease of American spearmint by Tobacco ring spot virus. Plant Dis Reptr 46:623–624 Tzanetakis IE, Postman JD, Martin RR (2005a) A member of the Closteroviridae from mint with similarities to all three genera of the family. Plant Dis 89:654–658 Tzanetakis IE, Postman JD, Martin RR (2005b) Characterization of a novel member of the family Closteroviridae from Mentha spp. Phytopathology 95:1043–1048 Tzanetakis IE, Postman JD, Martin RR (2006) Mint virus X: a novel potexvirus associated with symptoms in ‘Variegata’ mint. Arch Virol 151:143–153

M

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Mercurialis ambigua (Mercurial)

Tzanetakis IE, Postman JD, Martin RR (2007) Identification, detection and transmission of a new Vitivirus from Mentha. Arch Virol 152:2027–2033 Tzanetakis IE, Postman JD, Samad A, Martin RR (2010) Mint viruses: beauty, stealth, and disease. Plant Dis 94:4–12 Vicchi V, Bellardi MG (1988) The role of weeds in the epidemiology of gladiolus viruses and MLO. Acta Hortic 234:371–378 Zhou XG, Yuan XR, Wang SJ (1990) Two new virus diseases found on spearmint. Acta Agric Shanghai 6:45–52

Mercurialis ambigua (Mercurial) Synonyms Mercurialis annua Family: Euphorbiaceae

Weed host

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Mercurialis annua was reported from Italy (Grieco et al. 2000). The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sapinoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

Tomato yellow leaf curl virus Taxonomic position Genus: Begomovirus

(TYLCV)

Family: Geminiviridae

TYLCV infection in plants of Mercurialis ambigua was reported from southern Spain and Cyprus (Sanchez-Campos et al. 2000; Papayiannis et al. 2011). The virus-infected mercurial plants show abnormal upward leaf curling and leaf distortion symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. The virus is transmissible by grafting but not transmissible by contact between plants. For more details of TYLCV, refer to Solanum lycopersicum.

References Grieco PD, Conte D, Munno I, Nuzzaci M, de Stradis A (2000) Tomato spotted wilt virus (TSWV) on weeds and wild plants in Metapontino, Basilicata. Informatore Fitopatologico 50:43–46 Papayiannis L, Katis NI, Idris AM, Brown JK (2011) Identification of weed hosts of Tomato yellow leaf curl virus in Cyprus. Plant Dis 95:120–125 Sanchez-Campos S, Navas-Castillo J, Monci F, Diaz JA, Moriones E (2000) Mercurialis ambigua and Solanum luteum: two newly discovered natural hosts of tomato yellow leaf curl geminiviruses. Eur J Plant Pathol 106:391–394

Merremia spp.

1537

Merremia spp. Family: Convolvulaceae

Ornamental

Merremia mosaic Puerto Rico virus Taxonomic position Genus: Begomovirus

(MerMPRV)

Family: Geminiviridae

Geographical distribution MerMPRV infection in plants of Merremia spp. was reported from Puerto Rico (Idris and Brown unpublished - FJ944021; He et al. 2012). Symptoms and host(s) The virus-infected merremia plants exhibit mosaic symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate), incomplete icosahedra, and T = 1, 22  38 nm with a single coat protein. The genome is a bipartite consisting of two circular, single-stranded DNA components. DNA-A consists of 2640 nt (FJ944021 = NC_015490) and DNA-B of 2585 nt (FJ944022 = NC_015491) (Briddon 2001; Brown et al. 2015; Zerbini et al. 2017).

Merremia mosaic virus Taxonomic position Genus: Begomovirus

(MerMV)

Family: Geminiviridae

Geographical distribution MeRMV was reported in plants of Merremia quinquefolia from Puerto Rico (Bird et al. 1975; Brown et al. 2001). Symptoms and host(s) The virus-infected merremia plants exhibit bright yellow or green mosaic symptoms. The natural host range of this virus includes tomato, beans, and Datura stramonium. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate), incomplete icosahedra, and T = 1, 22  38 nm with a single coat protein. The genome is circular a single-stranded DNA and composed of two segments, DNA-A of

M

1538

Mikania micrantha (Bitter vine)

2557 nt (AF068636 = NC_007965) and DNA-B of 2492 nt (AY965899 = NC_007966) (Briddon 2001; Brown et al. 2001, 2015; Zerbini et al. 2017).

References Bird J, Sánchez J, Rodriguez RL, Julia FJ (1975) Rugaceous (whitefly-transmitted) viruses in Puerto Rico. In: Bird J, Maramorosch K (eds) Tropical diseases of legumes. Academic, London, pp 3–25 Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer Index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Idris AM, Torres-Jerez I, Banks GK, Wyatt SD (2001) The core region of the coat protein gene is highly useful for establishing the provisional identification and classification of begomoviruses. Arch Virol 146:1581–1598 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 He Z, Idris AM, Tang Y, Brown JK (2012) Infectivity and synergism for two monopartite begomoviruses and a bipartite begomovirus isolated from endemic Merremia species in Puerto Rico. www.apsnet.org/meetings/documents/2012_ meeting_abstracts/aps12ab0178.htm. Accessed 17 Apr 2016 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Mikania micrantha (Bitter vine) Family: Asteraceae

Gentian mosaic virus

Medicinal plant

(GeMV)

Synonyms Mikania micrantha wilt virus (MMWV) Taxonomic position Genus: Fabavirus

Family: Secoviridae

MMWV infection in plants of Mikania micrantha was reported from China (Wang et al. 2008, 2013). The virus-infected bitter vine plants exhibit wilting, crumpled, and malformed symptoms. The virus is transmitted by the aphid vector, Myzus persicae, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of GeMV, refer to Gentiana spp.

References Wang RL, Ding LW, Sun QY, Li J, Xu ZF, Peng SL (2008) Genome sequence and characterization of a new virus infecting Mikania micrantha HBK. Arch Virol 153:1765–1770 Wang R-L, Chen Y, Zhang H, Guan A-M, Wang Z-Y, Gong X, Yin Y, Zeng R-S (2013) Host range, transmitting vector, and damage characteristics of Mikania micrantha wilt virus. [In Chinese]. Chin J Ecol 32:72–77

Mimulus spp. (Monkey flower)

1539

Mimosa spp. (Touch-me-not) Family: Fabaceae

Weed host

Mimosa yellow leaf curl virus Taxonomic position Genus: Begomovirus

(MiYLCV)

Family: Geminiviridae

Geographical distribution MiYLCV infection in plants of Mimosa spp. was reported from Vietnam (Ha et al. 2008). Symptoms and host(s) The virus-infected touch-me-not plants exhibit yellow leaf curl symptoms. Transmission The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate), incomplete icosahedra, T = 1, 22  38 nm, with a single coat protein. The genome consists of a single molecule of circular, single-stranded DNA (DNA-A) of 2757 nt (DQ641695 = NC_009546), A betasatellite of 1358 nt (DQ641710) and DNA-1 of 1378 nt (DQ641719) are known to be associated with the virus (Briddon 2001; Ha et al. 2008; Brown et al. 2015; Zerbini et al. 2017).

References Briddon RW (2001) Begomovirus. Geminiviridae. In: The Springer Index of Viruses. Springer, New York, pp 340–348. https://doi.org/10.1007/3-540-31042-8_55 Brown JK, Zerbini FM, Navas-Castillo J, Moriones E, Ramos-Sobrinho R, Silva JC, Fiallo-Olivé E, Briddon RW, Hernández-Zepeda C, Idris A, Malathi VG, Martin DP, Rivera-Bustamante R, Ueda S, Varsani A (2015) Revision of Begomovirus taxonomy based on pairwise sequence comparisons. Arch Virol 160(6):1593–1619 Ha C, Coombs S, Revill P, Harding R, Vu M, Dale J (2008) Molecular characterization of begomoviruses and DNA satellites from Vietnam: additional evidence that the New World geminiviruses were present in the Old World prior to continental separation. J Gen Virol 89:312–326 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Mimulus spp. (Monkey flower) Family: Phrymaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Ornamental

(CMV)

Family: Bromoviridae

M

1540

Minthostachys mollis (Peperina)

CMV infection in plants of Mimulus guttatus was reported from the USA (Carr et al. 2003). Plants of monkey flower infected with CMV typically showed chlorosis and leaf curling, frequently accompanied with severe flower deformation. The virus is transmitted by a number of aphid vectors in a nonpersistent manner, and also by mechanical sap-inoculation to a large number of host plants. For more details of CMV, refer to Cucumis sativus.

Impatiens necrotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(INSV)

Family: Tospoviridae

Infection of Mimulus  hybridus by INSV has been reported from the USA (Daughtrey 2000). The virus-infected monkey flower plants exhibit necrotic spotting and ring-spot symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of INSV, refer to Impatiens spp.

References Carr DE, Murphy JF, Eubanks MD (2003) The susceptibility and response of inbred and outbred Mimulus guttatus to infection by Cucumber mosaic virus. Evol Ecol 17:85–103 Daughtrey M (2000) Diseases of mimulus, monkey-flower (Mimulus  hybridus hort. ex Siebert & Voss). http://www. apsnet.org/publications/commonnames/Pages/Mimulus.aspx

Minthostachys mollis (Peperina) Family: Lamiaceae

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

Medicinal

(CMV)

Family: Bromoviridae

CMV infection in plants of Minthostachys mollis was reported from Argentina (Rodriguez Pardina et al. 2013). The virus-infected peperina plants exhibit yellow mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of hosts. For more details of CMV, refer to Cucumis sativus.

References Rodriguez Pardina P, Ojeba M, Biderbost E, Di Feo L (2013) Detection and characterization of a Cucumber mosaic virus isolate infecting peperina, a species native to Argentina. Agriscientia 30:79–85

Mirabilis spp.

1541

Mirabilis spp. Family: Nyctaginaceae

Ornamental

Basella rugose mosaic virus Taxonomic position Genus: Potyvirus

(BaRMV)

Family: Potyviridae

BaRMV infection in plants of Mirabilis jalapa (four o’ clock) was reported from China (JN250997) (Wang et al. 2012). The virus-infected mirabilis plants exhibit mosaic and malformation symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sapinoculation. For more details of BaRMV, refer to Basella alba (Syn.) Basella rubra.

Chilli leaf curl India virus Taxonomic position Genus: Begomovirus

(ChiLCINV)

Family: Geminiviridae

ChiLCINV infection in plants of Mirabilis jalapa was reported from India (Jaidi et al. 2017). The virusinfected mirabilis plants exhibit upward leaf curling and stunting symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. For more details of ChiLCINV, refer to Capsicum annuum/Capsicum frutescens.

Mirabilis jalapa mottle virus Taxonomic position Genus: Carlavirus

(MjMV)

Family: Betaflexiviridae

Geographical distribution MjMV infection in plants of Mirabilis jalapa was reported from Texas, USA (Hatlestad et al. 2011). Symptoms and host(s) The virus-infected mirabilis plants exhibit slight leaf mottling and leaf wrinkling symptoms. Transmission The virus is mechanically sap-transmissible. Virion properties and genome The virions are flexuous filaments about 610–700 nm in length and 12–15 nm in diameter with helical symmetry. The genome consists of a single molecule of linear ssRNA of 8315 nt (JN039374 = NC_016080) and comprises six ORFs, encoding, in order, the replication-related proteins, the putative MPs (TGB), the CP, and a putative nucleic acid-binding regulatory protein. Coat protein subunits are of one type and 31–36 kDa in size (Adams et al. 2004).

M

1542

Mirabilis leaf curl virus Taxonomic position Genus: Begomovirus

Mirabilis spp.

(MirLCV)

Family: Geminiviridae

Geographical distribution MirLCV infection in plants of Mirabilis jalapa was reported from Northern India (Kulshreshtha et al. 2017). Symptoms and host(s) The virus-infected mirabilis plants exhibit severe leaf curling and mottling symptoms. Transmission The virus is transmitted by the whitefly vector Bemisia tabaci in a circulative, non-propagative manner. Virion properties and genome The virions are twinned (geminate) incomplete icosahedra, T = 1, 22 x 38 nm with a single coat protein. The genome is a monopartite circular ssDNA and consists of 2778 nt (LK054801 = NC_024491) and includes six ORFs, with two in the virion-sense strand (i.e., V1 and V2) and four in the complementarysense strand (i.e., C1–C4). Tomato leaf curl Patna betasatellite DNA molecule which consists of 1367 nt (LK054803) is associated with MLCV (Zhou 2013; Kulshreshtha et al. 2017; Zerbini et al. 2017).

Mirabilis mosaic virus Taxonomic position Genus: Caulimovirus

(MiMV)

Family: Caulimoviridae

Geographical distribution MiMV infection was first reported in plants of Mirabilis nyctaginea from Illinois, USA, by Brunt and Kitajima (1973). The virus spreads in the USA. Symptoms and host(s) The virus-infected mirabilis plants exhibit symptoms of chlorotic vein-banding of tip leaves and interveinal chlorosis of older leaves. Transmission The virus is transmitted by the aphid vector, Myzus persicae, in a semi-persistent manner. This virus is transmissible by mechanical sap-inoculation. The virus is not transmissible by contact between plants, not transmitted by seed, and not transmitted by pollen. Virion properties and genome The virions are isometric, non-enveloped, and 50 nm in diameter with no obvious surface structure. The genome is a monopartite, circular, double-stranded DNA of 7857 bp (AF454635 = NC_004036) (Donson and Hull 1983; Dey and Maiti 2002; Hohn 2011).

Mirabilis spp.

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Parietaria mottle virus Taxonomic position Genus: Ilarvirus

(PMoV)

Family: Bromoviridae

PMoV infection in plants of Mirabilis jalapa was reported from Italy (Parrella 2002). The virusinfected mirabilis plants showed symptoms of mild mosaic, leaf malformations of the upper leaves, and necrotic line patterns in some of the basal leaves. The virus is transmitted by thrips vectors, and the virus is also mechanically sap-transmissible. For more details of PMoV, refer to Parietaria officinalis.

Tobacco curly shoot virus Taxonomic position Genus: Begomovirus

(TbCSV)

Family: Geminiviridae

TbCSV infection in plants of Mirabilis jalapa was reported from Yunnan, China, by Xiong et al. (2010). The virus-infected mirabilis plants show typical yellow vein, leaf curling, and yellowing symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. For more details of TbCSV, refer to Nicotiana tabacum.

Tomato chlorotic spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(TCSV)

Family: Tospoviridae

TCSV infection in plants of Mirabilis jalapa was reported from Sao Paulo (Brazil) (Duarte et al. 2016). The virus-infected mirabilis plants exhibit systemic foliar mosaic symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation. For more details of TCSV, refer to Solanum lycopersicum.

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

TSWV infection in plants of Mirabilis japala was reported from Iran (Moini and Izadpanah 2000). The virus-infected mirabilis plants exhibit marginal necrosis of leaves and flower deformation and wilting symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Adams MJ, Antoniw JF, Bar-Joseph M, Brunt AA, Candresse T, Foster GD, Martelli GP, Milne RG, Fauquet CM (2004) The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. Arch Virol 149:1045–1060

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Miscanthus spp. (Silver grass)

Brunt AA, Kitajima EW (1973) Intracellular location and some properties of Mirabilis mosaic, a new member of the cauliflower mosaic group of viruses. Phytopathol Z 76:265–275 Dey N, Maiti IB (2002) Promoter deletion and comparative expression analysis of the Mirabilis mosaic caulimovirus (MMV) sub-genomic transcript (Sgt) promoter in transgenic plants. Transgenics 4:35–53 Donson J, Hull R (1983) Physical mapping and molecular cloning of Caulimovirus DNA. J Gen Virol 64:2281–2288 Duarte LML, Chaves ALR, Kitajima EW, Rodrigues LK, Harakava R, Alexandre MAV (2016) First report of Tomato chlorotic spot virus on Mirabilis jalapa. Aust Plant Dis Notes 11:6 Hatlestad GJ, Elam L, Gonzalez A, Lloyd AM (2011) Mirabilis jalapa mottle virus: a new carlavirus infecting four o’clocks. Arch Virol 156:2109–2111 Hohn T (2011) Caulimovirus. Caulimoviridae. In: The Springer Index of Viruses. Springer, New York, pp 271–277. https://doi.org/10.1007/978-0-387-95919-1_41 Jaidi M, Kumar S, Raj SK (2017) First report of Chilli leaf curl India virus infecting Mirabilis jalapa in India. New Dis Rep 35:2 Kulshreshtha A, Roshan P, Sharma D, Hallan V (2017) Molecular characterization of a new begomovirus infecting Mirabilis jalapa in Northern India. Arch Virol 162:2163–2167 Moini AA, Izadpanah K (2000) New hosts for Tomato spotted wilt virus in Tehran. Iran J Plant Pathol 36:104–105 Parrella G (2002) First report of Parietaria mottle virus in Mirabilis jalapa. Plant Pathol 51:401 Wang JG, Peng JJ, Chen HR, Chen SY (2012) First report of Basella rugose mosaic virus infecting four o’clock (Mirabilis jalapa) in China. Plant Dis 96:294 Xiong Y, Qing L, Ren F, Li F, Sun XC (2010) First report of Tobacco curly shoot virus on Mirabilis jalapa Linn. in China. J Plant Pathol 92:546 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133 Zhou X (2013) Advances in understanding begomovirus satellites. Annu Rev Phytopathol 51:357–381

Miscanthus spp. (Silver grass) Family: Poaceae

Forage crop

Barley yellow dwarf virus

(BYDV)

Taxonomic position BYDV is a tentative member of the genus Luteovirus and family Luteoviridae BYDV infection in plants of Miscanthus sacchariflorus was reported from the United Kingdom (Christian et al. 1994). The virus-infected silver grass plants exhibit symptoms of reddish-purple and discoloration on older leaves. The virus is transmitted by aphid vectors, Rhopalosiphum padi and Sitobion avenae, in a circulative, non-propagative manner; serological tests indicated the presence of BYDV-MAV, BYDV-PAV, and CYDV-RPV (as BYDV-RPV) in various combinations. No mechanical transmission of this virus is reported. For more details of BYDV-MAV, BYDV-PAV, and CYDV-RPV, refer to Hordeum vulgare.

Miscanthus streak virus Taxonomic position Genus: Mastrevirus

(MiSV)

Family: Geminiviridae

Miscanthus spp. (Silver grass)

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Geographical distribution MiSV infection was first reported in plants of Miscanthus sacchariflorus from Chiba, Japan, by Yamashita et al. (1979). The virus spreads in Japan (Yamashita et al. 1985). Symptoms and host(s) The virus-infected silver grass plants exhibit symptoms of chlorotic streaking and plant stunting. Transmission The virus is transmitted by leafhopper vectors in a persistent, circulative, and non-propagative manner. The virus is not transmissible by mechanical inoculation. Virion properties and genome The virions are twinned (geminate), incomplete icosahedra, T = 1, 22 x 38 nm, with a single coat protein. The genome is a monopartite circular, single-stranded DNA of 2672 nt (D01030 = NC_003379) (Chatani et al. 1991; Palmer and Rybicki 1998; Boulton and Davies 2011; Muhire et al. 2013; Zerbini et al. 2017).

Sorghum mosaic virus Taxonomic position Genus: Potyvirus

(SrMV)

Family: Potyviridae

SrMV infection in plants of Miscanthus sinensis was reported from the USA (Grisham et al., 2012). The virus-infected silver grass plants exhibit mosaic symptoms. The virus is transmitted by aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SrMV, refer to Sorghum bicolor.

Switchgrass mosaic virus

(SwMV)

Taxonomic position SwMV is a tentative member of the genus Marafivirus and family Tymoviridae SwMV infection in plants of Miscanthus spp. was reported from the USA (Agindotan et al. 2013). The virus-infected silver grass plants exhibit mosaic symptoms. The virus is transmitted by leafhopper vectors. For more details of SwMV, refer to Panicum virgatum.

References Agindotan B, Okanu N, Oladeinde A, Voigt T, Long S, Gray M, Bradley C (2013) Detection of Switchgrass mosaic virus in Miscanthus and other grasses. Can J Plant Pathol 35(1):81–86 Boulton MI, Davies JW (2011) Mastrevirus. Geminiviridae. In: The Springer Index of Viruses. Springer, New York, pp 597–603. https://doi.org/10.1007/978-0-387-95919-1_83 Chatani M, Matsumoto Y, Mizuta H, Ikegami M, Boulton MI, Davies JW (1991) The nucleotide sequence and genome structure of the geminivirus miscanthus streak virus. J Gen Virol 72(Pt 10):2325–2331 Christian DG, Lamptey JNL, Forde SMD, Plumb RT (1994) First report of barley yellow dwarf luteovirus on Miscanthus in the United Kingdom. Eur J Plant Pathol 100:167–170 Grisham MP, Maroon-Lango CJ, Hale AL (2012) First report of Sorghum mosaic virus causing mosaic in Miscanthus sinensis. Plant Dis 96:150

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Molinia caerulea (Purple moor grass)

Muhire B, Martin DP, Brown JK, Navas-Castillo J, Moriones E, Zerbini FM, Rivera-Bustamante R, Malathi VG, Briddon RW, Varsani A (2013) A genome-wide pairwise-identity-based proposal for the classification of viruses in the genus Mastrevirus (family Geminiviridae). Arch Virol 158(6):1411–1424 Palmer KE, Rybiscki EP (1998) The molecular biology of mastreviruses. Adv Virus Res 50:183–234 Yamashita S, Doi Y, Yora K (1979) Ann phytopath. Soc Jpn 45:128 Yamashita S, Nonaka N, Namba S, Doi Y, Yora K (1985) Miscanthus streak virus, a geminivirus in Miscanthus sacchariflorus. Ann Phytopathol Soc Jpn 51:582–590 Zerbini FM, Briddon RW, Idris A, Martin DP, Moriones E, Navas-Castillo J, Rivera-Bustamante R, Roumagnac P, Varsani A, ICTV Report Consortium (2017) ICTV virus taxonomy profile: Geminiviridae. J Gen Virol 98:131–133

Molinia caerulea (Purple moor grass) Family: Poaceae

Molinia streak virus

Forage crop

(MoSV)

Taxonomic position MoSV is a tentative member of the genus Panicovirus and family Tombusviridae Geographical distribution MoSV was described first in Germany, and it has been suggested that it may be distributed throughout central Europe (Huth et al. 1973–1974; Huth 1974). Symptoms and host(s) The virus-infected plants of purple moor grass display chlorotic-streaked spots plus a slight chlorosis of whole plants, delayed flowering, and occasional characteristic white spikes. Molinia caerulea is the only known natural host of this virus. Transmission The virus is mechanically sap-transmissible to certain graminaceous species (Huth et al. 1973–1974). Virion properties and genome The virions are isometric, 28 nm in diameter (Huth et al. 1973–1974).

References Huth W (1974) Molinia streak virus – a new virus on grasses. Mikrobiologija 11:195–196 Huth W, Paul HL, Querfurth G, Lesemann DL (1973–1974) Molinia streak virus: a virus with isometric particles from Molinia caerulea. Intervirology 2:345–351

Mollugo verticillata (Indian chickweed) Family: Molluginaceae

Weed host

Tomato spotted wilt orthotospovirus Taxonomic position Genus: Orthotospovirus

(TSWV)

Family: Tospoviridae

Momordica charantia (Bitter gourd/Bitter melon/Balsam apple/Balsam pear)

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TSWV infection in plants of Mollugo verticillata was reported from the USA (Groves et al. 2002). The virus-infected Indian chickweed plants exhibit yellow spots and ring-spot symptoms. The virus is transmitted by thrips vectors in a persistent-propagative manner, and also by mechanical sap-inoculation to a large number of herbaceous hosts. The virus is transmissible by grafting but not by contact between plants. For more details of TSWV, refer to Solanum lycopersicum.

References Groves RL, Walgenbach JF, Mayor JW, Kennedy GG (2002) The role of weed hosts and Tobacco thrips, Frankliniella fusca, in the epidemiology of Tomato spotted wilt virus. Plant Dis 86:573–582

Moluccella laevis (Bells-of-Ireland) Family: Lamiaceae

Weed host

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

CGMMV infection in plants of Moluccella laevis was reported from Israel (Shargil et al. 2017). The virus is transmissible by mechanical sap-inoculation, and also by contact between plants. For more details of CGMMV, refer to Cucumis sativus.

References Shargil D, Smith E, Lachman O, Reingold V, Darzi E, Tam Y, Dombrovsky A (2017) New weed hosts for Cucumber green mottle mosaic virus in wild Mediterranean vegetation. Eur J Plant Pathol 148:473–480

Momordica charantia (Bitter gourd/Bitter melon/Balsam apple/ Balsam pear) Family: Cucurbitaceae

Vegetable

Chayote yellow mosaic virus Taxonomic position Genus: Begomovirus

(ChaYMV)

Family: Geminiviridae

ChaYMV infection in plants of Momordica charantia was reported from Benin, Nigeria, and Togo (Leke et al. 2016). The virus-infected bitter gourd plants exhibit yellow mosaic symptoms. The virus is

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Momordica charantia (Bitter gourd/Bitter melon/Balsam apple/Balsam pear)

transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. Chayote yellow mosaic Benin betasatellite DNA molecule and consists of 1367 nt (KT454829) is associated with ChaYMV (Zhou 2013). For more details of ChaYMV, refer to Sechium edule.

Cucumber green mottle mosaic virus Taxonomic position Genus: Tobamovirus

(CGMMV)

Family: Virgaviridae

CGMMV infection in plants of Momordica charantia was reported from India (Pooja Kumari et al. 2016; Nagendran et al. 2017). The virus is not transmitted by any insect vector. The virus is transmitted through soil and irrigation water contaminated with infected plant debris. The virus is also pollen transmitted. For more details of CGMMV, refer to Cucumis sativus.

Cucumber mosaic virus Taxonomic position Genus: Cucumovirus

(CMV)

Family: Bromoviridae

CMV infection in plants of Momordica charantia was reported from northern Italy, Japan, China, and India (Kiranmai et al. 1998; Takami et al. 2006; Davino et al. 2012; Zhu et al. 2017). The virus-infected bitter gourd plants exhibit various degrees of mosaic mottling symptoms on the leaves; the young leaves are generally filiformed and have reduced size of the lamina. The virus is transmitted by a number of aphid vectors in a non-persistent manner, and also by mechanical sap-inoculation to a large number of host plants. For more details of CMV, refer to Cucumis sativus.

Cucurbit aphid-borne yellows virus Taxonomic position Genus: Polerovirus

(CABYV)

Family: Luteoviridae

CABYV infection in plants of Momordica charantia was reported from China, Tunisia, Vietnam, Indonesia, India, Sri Lanka, Thailand, Taiwan, Tunisia, Italy, France, Spain, and China (Xiang et al. 2008; Knierim et al. 2010; Suveditha et al. 2017). The virus-infected bitter gourd plants exhibit symptoms of interveinal mottling and yellowing of older leaves combined with thickening and brittleness. These were observed in several melon types, both under plastic tunnels and in the open field. Besides bitter gourd, other host crops include wax gourd, watermelon, cucumber, zucchini, squash, ridge gourd, snake gourd, and lettuce. The virus is transmitted by aphid vectors, Aphis gossypii and Myzus persicae, in a circulative, non-propagative manner, and not transmissible by mechanical sapinoculation. For more details of CABYV, refer to Cucumis melo.

Indian cassava mosaic virus Taxonomic position Genus: Begomovirus

(ICMV)

Family: Geminiviridae

Momordica charantia (Bitter gourd/Bitter melon/Balsam apple/Balsam pear)

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The association of Indian cassava mosaic virus (ICMV) with yellow mosaic disease of Momordica charantia was detected for the first time in Tamil Nadu, South India (Rajinimala and Rabindran 2007). The symptoms that characterize the yellow mosaic disease of bitter gourd are first visible on young leaves as mosaic and mottling only. The mottling usually starts at the edges of the leaf and advance inwards. Subsequently, chlorotic patches appear on leaves, and in advanced stages of infection, the entire leaf becomes chlorotic with a few, small patches of green tissue remaining over the leaf area. Blistering on leaves was also observed (Khan et al. 2002). The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. Through mechanical sap-inoculation, the virus was not transmitted through sap from bitter gourd to bitter gourd, but the virus was positively transmissible by sap to Nicotiana benthamiana, and symptoms of rosetting were observed. For more details of ICMV, refer to Manihot esculenta.

Melon yellow spot orthotospovirus Taxonomic position Genus: Orthotospovirus

(MYSV)

Family: Tospoviridae

MYSV infection of balsam pear (Momordica charantia) was found in Kochi Prefecture, Japan (Okuda et al. 2002; Takeuchi et al. 2009). The virus-infected balsam pear plants show necrotic spots and yellowing on leaves. The virus is transmitted by the thrips vector, Thrips palmi, in a persistentpropagative manner. For more details of MYSV, refer to Cucumis melo.

M Papaya ringspot virus Taxonomic position Genus: Potyvirus

(PRSV)

Family: Potyviridae

PRSV was reported to infect plants of Momordica charantia in India, China, Cuba, Jamaica, and the USA (Chin et al. 2007; Cabrera-Mederos et al. 2015; Kumari et al. 2016; Zhu et al. 2016). The virusinfected bitter gourd plants exhibited severe mosaic and deformations of leaves. The virus was saptransmissible from bitter gourd to Cucurbita moschata but not to papaya plants, indicating that it was the W pathotype of PRSV infecting bitter gourd in Cuba (Cabrera-Mederos et al. 2015), whereas PRVS-P was reported in Jamaica (Chin et al. 2007). The virus is transmitted by aphid vectors in a nonpersistent manner, and also by mechanical sap-inoculation. For more details of PRSV, refer to Carica papaya.

Pepper leaf curl Bangladesh virus Taxonomic position Genus: Begomovirus

(PepLCBV)

Family: Geminiviridae

PepLCBV infection in plants of Momordica charantia was reported from India (Raj et al. 2010). The virus-infected bitter gourd plants exhibit symptoms of a severe yellow mosaic and slight curling of leaf tissues. Infected plants had less fruit, which were smaller in size than those produced by healthy plants.

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Momordica charantia (Bitter gourd/Bitter melon/Balsam apple/Balsam pear)

The virus is transmitted by the whitefly vector, Bemisia tabaci, in a circulative, non-propagative manner. For more details of PepLCBV, refer to Capsicum annuum.

Squash vein yellowing virus Taxonomic position Genus: Ipomovirus

(SqVYV)

Family: Potyviridae

SqVYV infection in plants of Momordica charantia was reported from Puerto Rico (Adkins et al. 2008; Acevedo et al. 2013). The virus-infected bitter gourd plants exhibit vein-clearing symptoms. The virus is transmitted by the whitefly vector, Bemisia tabaci, in a non-persistent manner, and also by mechanical sap-inoculation. For more details of SqVYV, refer to Cucurbita pepo.

Tomato leaf curl New Delhi virus

(ToLCNDV)

Synonyms Bitter gourd yellow vein virus (BGYVV) Taxonomic position Genus: Begomovirus

Family: Geminiviridae

T