Mealybugs of California with Taxonomy, Biology and Control of North American Species (Homoptera: Coccoidea: Pseudococcidae) [Reprint 2020 ed.] 9780520338227

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Mealybugs of California WITH TAXONOMY, BIOLOGY AND CONTROL OF NORTH AMERICAN S P E C I E S

MEALYBUGS OF CALIFORNIA With Taxonomy, Biology, and Control of North American Species (Homoptera: Coccoidea: Pseudococcidae)

Howard L. McKenzie

UNIVERSITY OF CALIFORNIA PRESS

• BERKELEY AND LOS ANGELES 1967

The buckram embossment on the cover of this book depicts the Giant Mealybug, Phenacoccus giganteus McKenzie.

University of California Press Berkeley and Los Angeles, California Cambridge University Press London, England Copyright © 1967 by The Regents of the University of California Library of Congress Catalog Card Number: 67-10279 Printed in the United States of America

To the late PROFESSOR G. F. FERRIS and the late DR. H A R O L D MORRISON, eminent coccidologists for over forty years, and close friends who contributed so much to our knowledge of this family of insects.

CONTENTS

INTRODUCTION

1

Acknowledgments Scope of Study List of Colored Photographs

2 4 4

ECONOMIC IMPORTANCE

6

CONTROL MEASURES Control Organic Phosphates Chlorinated Hydrocarbons Ground Mealybug Control Miscellaneous Chemical Control Adversities Caused by Chemicals Cultural Methods

12 12 12 13 13 14 14 14

ECOLOGY ( Mealybug Habitats, Ant Associations )

16

Macro-environmental Preferences Micro-environmental Preferences Perennial-Flowering Plants (Excluding Grasses) Annual-Flowering Plants (Grasses Excluded) Grasses Conifers Ferns Inanimate Objects Multiple Host Positions Ant Associations Benefits to Mealybugs Benefits to Ants Strange Aspects of Ant-Mealybug Associations

16 16 16 17 17 18 19 19 19 20 20 22 22

BIOLOGY (Life History, Annual Successions) Life History Oviposition First Instar Nymph Second Instar Nymph Third Instar Nymph Adult Female Fourth Instar Male Fifth Instar Male (Adult) Parthenogenesis Instars and Generations Successions in Mealybugs

24 24 24 24 24 24 24 25 25 27 27 27

viii

CONTENTS CYTOLOGY Genetic Systems Parthenogenesis Sex Determination Chromosome Number Technique F I E L D AND LABORATORY METHODS Collecting Methods Choosing a Location Collecting on Foliage and Stems Collecting on Branches Collecting on Crown and Roots Collecting on Grass Collecting on Conifers Preserve Plant Material Preparation of Slide Mounts Temporary Mounts Permanent Mounts MORPHOLOGY AND CLASSIFICATION Morphology Segmentation Head Thorax Abdomen Pores and Ducts Body Setae Classification (Definition of Family Pseudococcidae) N O R T H AMERICAN GENERA O F PSEUDOCOCCIDAE Field Characteristics of Mealybug Genera Key to Genera of North American Pseudococcidae TAXONOMY O F SPECIES (Keys to North American Species) List of California Mealybugs

28 28 29 29 30 30 32 32 32 32 33 33 33 33 33 34 34 34 36 36 36 36 38 39 39 40 40 41 42 44 48 48

DISTRIBUTION TABLE

493

LITERATURE CITED

499

HOST INDEX (With Associated Mealybugs)

501

G E N E R A L INDEX

518

ADDENDUM

526

INTRODUCTION

Mealybugs1 are slow moving insects which, in the female stage, are characteristically elongate-oval, soft, and with distinct segmentation. They are usually thinly or thickly covered with a mealy or cottony wax secretion which often extends from along the sides in a series of short filaments, and with usually two longer ones at the posterior end of body (see kodachrome prints and water color paintings in this book). Their mouthparts are thread like, often longer than the body itself, inserted through the tough tissue or bark to suck juices from the host; and they are among the most serious pests of plant life. They vary in size from very minute young, 0.50 mm, to mature adult females measuring as much as 8.00 mm, or perhaps 9.00 mm, in length. The eggs are usually laid in loose masses of cottony wax or felt-like ovisacs. Some species bear living young. The males, seldom seen, are delicate, winged or infrequently apterous (wingless), possessing two conspicuous long caudal wax filaments. They have abortive mouth parts and take no food. Identifications of mealybugs in practically all cases are based upon the adult female, since the males are so rarely available. Consequently, in this study the males are excluded from consideration. Historically, most of the literature on California mealybugs is confined to a few isolated publications. The first article pertaining to this subject was written by Carnes (1907) and entitled "The Coccidae of California." It was published in the second biennial report for 1905-1906, of the California State Commissioner Horticulture. In this publication twentynine (29) mealybugs species were listed which had been recorded, in the entomological literature of the world, as occurring in California. One (1) of these species had been taken in quarantine, but not established in the State. Eleven (11) of the more important pseudococcid species on the list were described in considerable detail, and two (2) of them 1

were illustrated in ink to show gross external features. Later, Essig (1914) wrote an article entitled "The Mealybugs of California," published in The Monthly Bulletin, State Commission of Horticulture, Sacramento, California. In Essig's article an attempt was made to bring together all the information about this group of insects, with particular reference to those of economic importance. Although special attention was given to the economic forms, mention was also made of the noneconomic species in order that the reader would have a more comprehensive idea of the group as a whole. Ten (10) species were listed by Essig in the noneconomic group of mealybugs; eight (8) species were designated as economic on ornamental plants; while four (4) species were assigned as economic on cultivated plants and fruit trees. Two (2) other species were discussed, but were not classified as either economic or noneconomic. Except for certain photographs which fairly adequately depict certain California mealybugs in life, no detailed illustrations showing specific morphological pseudococcid structures accompanied this publication. In 1918, Ferris published a much more sophisticated treatment of the subject, titled The California species of mealybugs, printed by the Stanford University Press. This study included some 44 species for the state. Rather precise pen and ink drawings of the penultimate and anal lobe cerarii on the last two abdominal segments of the mealybug were included in this study. Prior to this more comprehensive treatment of California mealybugs, little had been done to contribute to an accurate identification of species involved. Ferris (1917, p. 5) stated that "of the nearly 100 species of mealybugs thus far described from North America, including some 35 from California, not more than three or four are recognizable at all on the basis of the published descriptions if taken apart from their typical host and their type locality."

Homoptera; Coccoidea; Pseudococcidae.

1

2

1NTROD

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Until Ferris' 1918 publication, the additional claim could safely be made that even under most favorable conditions, the identifications of many of our California species could not be accomplished with reasonable accuracy. In 1950, and again in 1953, Ferris produced the splendid volumes of his Atlas of the scale insects of North America, Series V and Volume VI, the Pseudococcidae (Parts I and II respectively), published by the Stanford University Press. In these volumes there are approximately 250 specific names available for North American species of pseudococcids, these including an indeterminate number of synonyms. Of this number some 89 species are recorded in the Atlas as occurring in California. All the species in the Atlas were magnificently illustrated. Ferris implied that the primary intention of this work was to clarify existing names, rather than to describe new species. In fact, he (1950, p. I l l ) stated, "no search has been made in the available material for new species and much material has deliberately been set aside and omitted from consideration. Only here and there have previously undescribed species been included and these only because they have been involved in the synonymy of other species through misidentification or because they assist in rounding out some group that would be inadequately considered if they were omitted." In 1960 I published an article on "Taxonomic study of California mealybugs, with descriptions of new species," this appearing in Hilgardia, a University of California Agricultural Sciences Publication. The purpose of this study was to describe and delineate new California species of mealybugs, and to comment briefly on the taxonomy of other named forms if necessary. A prediction was made at the time that additional undescribed pseudococcid species in the state were a practical certainty and that subsequent collections, particularly of those forms occurring underground, would no doubt reveal many of the new species which would have to be dealt with later on. Furthermore, several new species collected outside the state were described and included in this study to broaden our concepts of this important family of Coccoidea. It was planned that the material made available in this and four subsequent studies (McKenzie, 1961, 1962, 1964, and 1965), all published in the Hilgardia series, would ultimately be incorporated in a systematic exposition on Mealybugs of California, to appear as a companion piece to my publication (1956), The Armored Scale Insects of California. The Hilgardia series of five studies contains descriptions of 57 new species of mealybugs, including those described in other Western States and subsequently found in California. ACKNOWLEDGMENTS

I am particularly indebted to Dr. R. M. Bohart, Chairman, Department of Entomology, Davis, for suggesting that a systematic exposition on the mealybugs of California be undertaken as a companion

piece to my (1956) publication on Armored Scale Insects of California. Dr. Bohart has also read the manuscript of this book, offering helpful suggestions, especially in the area of the more technical taxonomic problems. At the outset of these investigations, both the late Professor G. F. Ferris, Stanford University, California, and Dr. Harold Morrison, United States Department of Agriculture, Washington, D. C., offered their wholehearted support of the project. In addition, Dr. Morrison made available distribution and host records of California pseudococcids represented in the United States National Collection of Coccodea (now the United States National Museum). It is difficult to express how very grateful I am to both Professor Ferris and Dr. Morrison for all they have done, and I only hope that they were well aware of this attitude long before they passed on. In 1960, officials at Stanford University presented as a gift to the University of California, Department of Entomology, Davis, the entire Stanford Coccoidea Collection accumulated over a period of 40 years by Professor Ferris. Without this magnificent collection to rely on, the present volume would have been infinitely more difficult to complete. Appreciation is again expressed to Stanford University for this splendid gift. In addition, permission has been granted by Stanford University Press for the use of many of Professor Ferris' original illustrations from his Atlas, Series V (1950), and Volume VI (1953). These drawings were placed in my custody at the time the Scale Insect Collection and Library were transferred to' the University of California at Davis (see pages 3 and 4 for the list of illustrations used). Mr. R. F. Wilkey, taxonomist, Bureau of Entomology, California Department of Agriculture, Sacramento, has been particularly co-operative in assisting me in these investigations, not only in preparing the necessary slide mounts, but also in helping to "screen out" and determine the various lots of field collections. The numberous pseudococcid slide preparations he has made for this study have contributed much, because of their excellence, to the accuracy of the final technical illustrations of the involved species. I hereby acknowledge his most willing assistance. In addition, the California Bureau of Entomolgy has maintained extensive and accurate collection records of mealybugs taken in California, and these data have been extensively used in this publication, with the approval of Mr. R. M. Harper, Bureau Chief. Mr. Douglass R. Miller, Research Assistant and Graduate Student, University of California, Department of Entomology, Davis has been responsible for collecting the majority of the new mealybug species encountered in these studies. In addition, he has assisted in preparing some of the chapters in this book, as well as some of the drawings depicting certain undescribed species, and has helped in proofreading much of the manuscript. I extend to him my sincere appreciation for his untiring assistance during these investigations.

1NTROD The chapter on cytology was prepared in collaboration with Dr. Spencer W. Brown, Geneticist, University of California, Berkeley. Actually, Dr. Brown used only a very small part of my material on parthenogenesis in the cytology chapter and, therefore, most of the information on this subject should be accredited to him. Sincere appreciation for his contribution is here indicated. Drs. Robert L. Usinger, Chairman, Department of Entomology, Berkeley, E. Gorton Linsley, Dean, College of Agriculture, Berkeley, and Alfred M. Boyce, Dean, College of Agriculture, Riverside, examined and unanimously approved the publication of these investigations by the University of California Press. Their support of these studies is most gratefully acknowledged. Dr. Maurice L. Peterson, University Dean of Agriculture and Director of the Agriculture Experiment Station, Berkeley, kindly made funds available to cover the costs of printing the watercolor paintings and kodachrome prints of various mealybugs included in this book. To Dr. Peterson, and to Dr. James H. Meyer, Dean, Agriculture, Davis, who submitted the original request asking that these colored plates be included in the book, I extend my deepest thanks. Dr. W. Harry Lange, Department of Entomology, Davis, assisted in taking many of the kodachrome illustrations presented herein. In addition, both Drs. Lange and E. M. Stafford, same department, read the chapter on Control Measures and offered helpful suggestions with particular reference to insecticidal terminology. Dr. O. G. Bacon, same department, has carefully read all the chapters of this book exclusive of the taxonomy section. I gratefully acknowledge this kind assistance. Dr. L. R. Brown, Department of Entomology, Riverside, and Mr. D. J. Bingham, Nursery Service, California Department of Agriculture, Sacramento, loaned several excellent kodachrome slides of certain mealybug species included in the colored plates of this book. I express my appreciation to them for the loan of these slides. Thanks are extended to Mr. L. E. Myers, Supervising Inspector, Los Angeles County Agricultural Commissioner's Office, who loaned me his extensive mealybug collection for use during these investigations. The Museum of the California Academy of Sciences, San Francisco, California, through the generosity of Dr. E. S. Ross, kindly loaned me the mealybug collection of the late Professor E. O. Essig which is deposited at that institution. This collection has proven especially helpful in this study because of Professor Essig's early work on California pseudococcids. The University of California, Department of Entomology, Riverside, recently turned over their entire collection of Coccoidea for deposition in the Museum, University of California, at Davis. I am much indebted to Mr. Harold Compere, Drs. S. E. Flanders and C. A. Fleschner, for this generous gift.

UCTION

3

Much valuable information relative to California mealybugs has come from this collection. Miss June McCaskill, Department of Botany, Davis, is responsible for many of the wildflower and shrub identifications, and Dr. Beecher Crampton, Department of Agronomy, Davis, determined numerous grasses (Gramineae) mentioned in this book. I am deeply indebted to these individuals for this kind assistance. Since January, 1962, the National Science Foundation, Washington, D. C., has generously provided funds for a grant supporting this study. Continued support of the project is contemplated for all of 1966, and possibly even beyond that time. I express my appreciation to this organization for the grant, which has promoted constructive research on this important subject. Several delineators have assisted in the preparation of certain pen and ink illustrations, and their initials appear individually, or in pairs on some of them. These individuals are: Mrs. Julia Z. litis, Mrs. Helen Court, Mrs. Mary Foley Benson, and Douglass R. Miller. I sincerely appreciate their help, and especially their concern for this work. Without doubt, they have contributed much to the accuracy of these studies. Special mention is made of the impressive watercolor paintings of mealybugs reproduced in this book. The artist, Mrs. Mary Foley Benson, is known to many entomologists because of the paintings she prepared of certain economic insects which appeared in the United States Department of Agriculture Yearbook for 1952. Mrs. Benson was formerly Chief Scientific Illustrator for the United States Department of Agriculture. Later she established a studio in Hollywood, California, where she prepared special illustrations for display and sale, as well as other commercial and fine art. Her work has been shown in many art galleries, a few of which include United States National Museum, and Lausburgh, Washington, D. C.; Carnegie Institute of Technology, and Hunt Botanical Library, Pittsburgh, Pennsylvania; Hollywood Association of Artists, and Frances Webb Galleries, Los Angeles, California. Permission has been granted by Stanford University Press, Stanford University, California, to use numerous illustrations included in G. F. Ferris' (1950 and 1953), Atlas of the Scale Insects of North America, Series V, and Volume VI, The Pseudococcidae (Parts I and I I ) . Certain species drawings have been slightly altered to include corrections, and these are followed by the symbol ( A ) . The original drawings of figures 64 and 120 were lost. Copies of these illustrations were, however, made from Ferris' Atlas, and these are followed by the symbol ( C ) . The following list records the species by figure numbers as they occur in the present book: Fig. 5. Amonostherium lichtensioides (A); Fig. 7. Anisococcus adenostomae; Fig. 8. A. crawii (A); Fig. 9. A. didymus (A); Fig. 10. A. ephedrae; Fig. 12. A.

4

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UCTION

oregonensis; Fig. 13. A. quercus ( A ) ; Fig. 15. Antonina crawii; Fig. 16. A. graminis; Fig. 17. A. pretiosa; Fig. 18. Cataenococcus olivaceus (A); Fig. 20. C. villosa; Fig. 32. Chorizococcus reducta ( A ) ; Fig. 33. C. rostellum; Fig. 39. Crisicoccus azaleae; Fig. 40. C. pini ( A ) ; Fig. 41. Cryptoripersia salina ( A ) ; Fig. 43. Cucullococcus vaccinii; Fig. 44. Discococcus caricis; Fig. 45. D. flabellatus ( A ) ; Fig. 46. D. graminis; Fig. 47. D. hopi ( A ) ; Fig. 48. D. simplex (A); Fig. 51. Distichlicoccus californicus (A); Fig. 53. D. salinus ( A ) ; Fig. 54. Dysmicoccus aciculus ( A ) ; Fig. 58. D. quercicolus ( A ) ; Fig. 60. D. ryani (A); Fig. 61. D. timberlakei ( A ) ; Fig. 63. Ehrhornia cupressi; Fig. 64. Ferrisia virgata (CA); Fig. 67. Heliococcus stachyos ( A ) ; Fig. 68. Heterococcus arenae ( A ) ; Fig. 69. H. pulverarius ( A ) ; Fig. 70. Humococcus atriplicis ( A ) ; Fig. 73. H. hilariae ( A ) ; Fig. 75. Hypogeococcus spinosus ( A ) ; Fig. 76. Misericoccus arenarius (A); Fig. 77. Nipaecoccus aurilanatus (A); Fig. 78. N. nipae (A); Fig. 79. Paludicoccus distichlium; Fig. 82. Phenacoccus artemisiae (A); Fig. 85. P. celtisifoliae ( A ) ; Fig. 86. P. colemani (A); Fig. 88. P. dearnessi; Fig. 89. P. defectus; Fig. 93. P. eremicus (A); Fig. 94. P. eriogoni ( A ) ; Fig. 98. P. gossypii ( A ) ; Fig. 101. P. helianthi ( A ) ; Fig. 104. P. lycii; Fig. 106. P. minimus; Fig. 108. P. pauperatus (A); Fig. 109. P. solani (A); Fig. 110. P. solenopsis ( A ) ; Fig. 111. Planococcus citri ( A ) ; Fig. 112. P. kraunhiae ( A ) ; Fig. 113. Pseudantonina arundinariae (A); Fig. 117. Pseudococcus fragilis (A); Fig. 119. P. longisetosus (A); Fig. 120. P. longispinus (CA); Fig. 129. Puto ambiguus; Fig. 130. P. arctostaphyli ( A ) ; Fig. 132. P. bryanthi; Fig. 134. P. cupressi (A); Fig. 145. P. yuccae ( A ) ; Fig. 147. Radicoccus kelloggi ( A ) ; Fig. 152. Rhizoecus cacticans; Fig. 153. R. californicus; Fig. 154. R. cyperalis; Fig. 157. R. falcifer; Fig. 159. R. graminis; Fig. 160. R. kondonis; Fig. 161. R. leucosomus; Fig. 167. Spilococcus andersoni; Fig. 168. S. atriplicis; Fig. 172. S. eriogoni ( A ) ; Fig. 174. S. gutierreziae ( A ) ; Fig. 175. S. implicatus (A); Fig. 177. S. larreae ( A ) ; Fig. 181. S. prosopidis (A); Fig. 183. S. sequoiae ( A ) ; Fig. 185. Stemmatomerinx decorata; Fig. 186. Tridiscus distichlii; Fig. 187. T. multiorbis (A); Fig. 188. Trionymus americanus ( A ) ; Fig. 189. T. caricis; Fig. 190. T. diminutus (A); Fig. 191. T. dolus ( A ) ; Fig. 192. T. festucae; Fig. 196. T. mocus ( A ) ; Fig. 197. T. modocensis ( A ) ; and Fig. 200. T. smithii (A). SCOPE OF STUDY

I have tried to present pertinent information about all known mealybug species found in California— 193 species in all. Of this number, 88 species have been named by the author, 32 of which are described as new in the present volume. In addition, the number of pseudococcid genera represented in California totals 35, and of this number, four (4) have been previously described by the author. Some higher-group names have been suggested for certain pseudococcid genera in North America. A complete bibliographical record is included, with reference to the original description of each genus and species; their type locality and host; North America distribution; additional hosts; external features and habitat; recognition characters (morphological structures as

viewed through microscope); notes about relationship to other species; and California distribution, with an accompanying outline map showing range in the state. Dichotomous keys to North American genera and species are presented. A common name has been assigned to each California species of mealybug. This study also contains information relative to the economic importance of mealybugs in California; a résumé of control measures as gleaned through a search of the literature; ecology (habitats, ant associations); biology (life history, annual successions); cytology; techniques used in collecting mealybugs and preparing them for critical study on microscope slides; and a detailed account of external morphology of the Pseudococcidae. Each included species is carefully and completely illustrated in an attempt to fill the needs of the systematic entomologist, the technician, and those specifically engaged in biological and chemical control of mealybugs. A distribution table is presented primarily for the benefit of the economic entomologist. A general index covering all phases of this study is included at the end of this volume. Twenty-one full pages of water color paintings, and four full pages of composite Kodachrome prints, including thirty-six exposures, of certain California mealybugs are provided in this work. It is hoped that these illustrations will enable entomologists, agriculturalists, nursery men, field workers, and home gardeners to recognize our more important mealybugs. LIST OF COLORED PHOTOGRAPHS

Many of the specimens used to produce the accompanying photographs were collected by various individuals at my request. Some of the photographs were taken by other people and subsequently turned over to me for use in this book. Dr. Leland R. Brown, Entomologist, University of California, Riverside Campus, made available ten of the photographs here used. Mr. David J. Bingham, California State Department of Agriculture, Nursery Service, Sacramento, and Mr. R. F. Wilkey, same department, Bureau of Entomology, kindly turned over to me four Kodachrome slides, and prints from these appear on the following pages. Dr. W. H. Lange, Entomologist, University of California, Department of Entomology, Davis Campus, gave specific instructions necessary to produce many of the accompanying colored photographs. To these individuals, who have so generously participated in this project, acknowledgments are here made. Most of the colored photographs are enlargements of the insects, and range from approximately 3.5 to 12.5X magnifications. It is hoped that these photographs will help in identifying at least a few of our more common California mealybugs. The species have not been arranged chronologically, but instead are assembled more for the purpose of eye appeal and beauty.

INTRODUCTION T h e following list includes specific collection data for each colored photograph: Color Plate I Planococcus citri (Risso), Davis, Yolo Co., California, February 23, 1960, on Coleus sp. (coleus) (H. L. McKenzie). Heterococcus pulverarius (Newstead), 8 miles east Tulelake, Modoc Co., California, May 31, 1961, on Poa sp. (bluegrass), (K. G. Baghott). Chorizococcus lounsburyi (Brain), Pacoima, Los Angeles Co., California, April 6, 1953, on Gasteria linqua (succulent) (L. R. Brown). Phenacoccus colemani Ehrhorn, 10.7 miles north Hat Creek, Shasta Co., California, May 23, 1963, under volcanic lava rock, associated with ants (T. R. Haig). Phenacoccus solani Ferris, Davis (Putah Creek), Yolo Co., California, November 10, 1960, on Compositae (roots) ( M . E . I r w i n ) . Pseudococcus obscurus Essig, Castroville, Monterey Co., California, August 31, 1962, on Solidago sp. (golden rod) (W. H. Lange). Phenacoccus gossypii Townsend and Cockerell, Davis Campus, Yolo Co., California, September 7, 1961, on Encelia farinosa (incienso) (H. L. McKenzie). Phenacoccus gossypii Townsend and Cockerell, Exeter, Tulare Co., California, October 4, 1962, on Lantana sp. (lantana) (W. R. Clark) (ovisacs). Phenacoccus gossypii Townsend and Cockerell, Davis (Campus), Yolo Co., California, September 7, 1961, on Encelia farinosa (incienso) (H. L. McKenzie) (ovisacs and immatures). Color Plate II Anisococcus adenostomae Ferris, 11 miles east Capella, Mendocino Co., California, May 28, 1959, on Adenostoma fasciculatum (chamise) (T. R. Haig). Spilococcus cactearum McKenzie, Berkeley (U. C. Botanical Gardens), Alameda Co., California, November 4, 1958, on Hamatocactus setispinus (cactus) (A. E. Pritchard and H. L. McKenzie). Spilococcus cactearum McKenzie, Berkeley (U. C. Botanical Gardens), Alameda Co., California, January 20, 1959, on Echinomastus johnsonii (cactus) (H. L. McKenzie) (ovisacs). Phenacoccus eschscholtziae McKenzie, 2 miles west Yucca Valley, San Bernardino Co., California, April 9, 1963, on Malacothrix califomica var. glabrata (D. R. Miller). Cryptoripersia salina (Ehrhorn), 2 miles south Kneeland, Humboldt Co., California, June 16, 1960, on Danthonia californica (oat grass) (K. Miller and T. R. Haig). Pseudococcus obscurus Essig, (species with lateral filaments) and Spilococcus cactearum McKenzie, Berkeley (U. C. Botanical Gardens), Alameda Co., California, November 4, 1958, on Hamatocactus setispinus (cactus) (H. L. McKenzie and A. E. Pritchard). Nipaecoccus nipae (Maskell), Coronado Island, San Diego Co., California, March 6, 1952, on Kentia sp. (palm) (C. O. Eads and L. R. Brown). Phenacoccus giganteus McKenzie, 12 miles south Shoshone, Inyo Co., California, April 13, 1963, on Franseria dumosa (burro-weed) (D. R. Miller). Phenacoccus giganteus McKenzie, 12 miles south Shoshone, Inyo Co., California, April 13, 1963, on Franseria dumosa (burro-weed) (D. R. Miller) (ovisac).

5

Color Plate III Phenacoccus graminosus McKenzie, Napa, Napa Co., California, January 1, 1959, on Lolium sp. (rye grass —sheaths above crown) (overwintering adult females) (H. L. McKenzie and R. P. Allen). Phenacoccus celtisifoliae (Hollinger), Dunnigan, Yolo Co., California, July 28, 1964, on Photinia arbutifolia (Christmas berry, toyon), (T. Kono and H. H. Keifer). Nipaecoccus aurilanatus (Maskell), Beverly Hills, Los Angeles Co., California, January 7, 1958, on Araucaria excelsa (Norfolk Island pine) (D. Farrell). Cataenococcus olivaceous (Cockerell), Lawndale, Los Angeles Co., California, January 24, 1961, on Yucca sp. (Spanish bayonet) (L. E. Myers). Rhizoecus pritchardi McKenzie, Berkeley, Alameda Co., California, November 4, 1958, on Adiantum sp. (maidenhair fern, roots) (H. L. McKenzie and A. E. Pritchard). Spilococcus implicatus Ferris, % mile north Kneeland, Humboldt Co., California, August 14, 1963, on Sequoia sempervirens (redwood) (T. R. Haig). Pseudococcus fragilis Brain, Martinez, Contra Costa Co., California, April 8, 1955, on Pittosporum tobira (Japanese pittosporum) (D. J. Bingham and Danielson). Rhizoecus kondonis Kuwana, Healdsburg, Sonoma Co., California, January 17, 1962, on Prunus domestica (prune roots) (W. W. Wiard and H. McCracken). Rhizoecus kondonis Kuwana, November, 1961, damage to Medicago sativa (alfalfa) (H. L. McKenzie and V. E. Burton). Color Plate IV Chnaurococcus trifolii (Forbes), Truckee, Nevada Co., California, August 26, 1965, on Gramineae (grass) (T. R. Haig). Anisococcus quercus (Ehrhorn), 8 miles east Alturas, Modoc Co., California, August 29, 1963, on Purshia tridentata (antelope brush) (R. M. Hawthorne and R. C. Hall). Ferrisia virgata (Cockerell), Brawley, Imperial Co., California, December 31, 1963, on Lantana sp. (lantana) (D. R. Miller). Scaptococcus milleri McKenzie, new species, 8 miles west Glamis, Imperial Co., California, in sand dunes on Eriogonum deserticola (buckwheat family) (D. R. and J. F. Miller). Pseudococcus maritimus (Ehrhorn), Bodega Bay, Sonoma Co., California, August 4, 1963, on Eriogonum latifolium (D. R. Miller). Anisococcus oregonensis Ferris, 3 miles northwest Spring Garden, Plumas Co., California, July 7, 1965, on Eriogonum umbellatum polyanthum (sulphur flower) (T. R. Haig). Puto californicus McKenzie, new species, 5 miles north San Bernardino, San Bernardino Co., California, July 23, 1963, on Lotus sp. (D. R. Miller). Rhizoecus cacticans Hambleton, Berkeley (U. C. Botanical Gardens), Alameda Co., California, November 4, 1958, on Hamatocactus sp. (cactus roots) (H. L. McKenzie). Puto acirculus McKenzie (immature), Santa Paula, Ventura Co., California, May 22, 1958, on Elymus condensatus (giant rye grass) (L. R. Gillogly and J. L.Weber).

ECONOMIC IMPORTANCE

In 1927, Harold Compere found in Australia two species of hymenopterous parasites, Coccophagus gurneyi Compere and Tetracnemus pretiosus Timberlake, which not only attacked the citrophilus mealybug, but were also capable of maintaining themselves throughout the winter months (Compere and Smith, 1932). These parasites gave almost complete control over this mealybug on all of its host plants (Ebeling, 1959). Planococcus citri (Risso), the citrus mealybug (col. pis. I and X I V ) , has been a sporadic pest in the United States since 1879 (Quayle, 1938). In California it is a pest in nearly all greenhouses and nurseries where it attacks a wide range of ornamental hosts. It is considered a serious pest of citrus in many parts of the world. In California it was for years rated as the most important citrus-infesting species until the introduction of the devastating citrophilus mealybug. Heavy infestations on citrus have never been widespread throughout all of the citrus groves, but have been localized. Damage caused by citrus mealybug is of the normal fruit contamination, honeydew, and sooty-mold type. In addition, at least in some strains, this mealybug has an apparent toxic effect on the plant host. On Coleus this damage is usually noted when the medial area along the midrib of the leaves (normally the area of heaviest infestation) turns brown. Chemical control of this insect is amazingly difficult, with parathion giving the best results. In biological control programs, partial control has been attained by a coccinellid predator Cryptolaemus montrouzieri (Mulsant) and a hymenopterous parasite, Leptomastidea abnormis (Girault) (De Bach and Schlinger, 1964). Pseudococcus maritimus (Ehrhorn), grape mealybug (col. pi. I V ) , is another species important primarily to grapes and pears in California. This pseudococcid has for some time been confused with another species, Pseudococcus obscurus Essig, and because of this has caused much confusion in the lit-

Mealybugs in California are serious plant pests, and as a result have caused considerable research in the fields of chemical and biological control. One of the most important California pseudococcid pests over the past half century has been the citrophilus mealybug, Pseudococcus fragilis Brain (col. pis. I l l and X V I ) . This species was first found in California in 1913 near Upland (Woglum, 1922) and rapidly became a serious pest of citrus to the point that it threatened the industry. An attempt to eradicate the species in this area at that time failed, and it gradually spread along the coast into Los Angeles, Orange, and Ventura Counties. Since that time it has been primarily restricted to the cooler, more humid regions. Injury to host, as is typical with most mealybug species, is caused by extracting plant sap, excreting honeydew which provides a medium for sooty-mold fungus, and production of a cottony wax material on the fruit and leaves. The excretion of sticky honeydew on the host, and the subsequent build up of sooty mold growing in this medium', create an unsightly appearance especially to the fruits, thus causing serious economic loss to the grower. Chemical control measures directed against the citrophilus mealybug were ineffectual. Biological control, on the other hand, was quite successful. This was first effected by a lady-beetle predator, Cryptolaemus montrouzieri (Mulsant), introduced by Koebele from Australia in 1892. The beetle, in both the adult and larval stages, successfully attacked the mealybugs, but because of the beetle's high mortality during the winter months in California, it was unable to maintain sufficient numbers to counteract the rapid spring increase in the mealybug population. A successful method of mass rearing of the beetle in insectaries was developed, and this proved to be an effectual means of controlling the pest. The mass rearing and liberation of cryptolaemus, however, constituted an annual program involving considerable expense. 6

SSi

COLOR PLATE I. Top row. LEFT: Plancoccus citri (Risso), citrus mealybug; CENHeterococcus pulverarius (Newstead), bluegrass mealybug; RIGHT: Chorizococcus lounsburyi (Brain), lily bulb mealybug. Middle row. LEFT: Phenacoccus colemani Ehrhorn, Coleman mealybug; CENTER: Phenacoccus solani Ferris, solanum mealybug; RIGHT: Pseudococcus obscurus Essig, obscure mealybug. Bottom row. LEFT: Phenacoccus gossypii Townsend and Cockerell, Mexican mealybug; CENTER: ovisac of same; RIGHT: ovisac and immature of same. TER:

COLOR PLATE II. Top row. LEFT: Anisococcus adenostomae Ferris, chamise mealybug; CENTER: Spilococcus cactearum McKenzie, cactus mealybug; RIGHT: white ovisacs of same among cactus spines. Middle row. LEFT: Phenacoccus eschscholtziae McKenzie, California poppy mealybug; CENTER: Cryptoripersia salina (Ehrhorn), felt sac mealybug; RIGHT: Pseudococcus obscurus Essig, obscure mealybug (large species with conspicuous lateral and caudal wax filaments), and Spilococcus cactearum McKenzie, cactus mealybug. Bottom row. LEFT: Nipaecoccus nipae (Maskell), coconut mealybug; CENTER: Phenacoccus giganteas McKenzie, giant mealybug; RIGHT: ovisac of same.

COLOR PLATE III. Top row. LEFT: Phenacoccus graminosus McKenzie, ryegrass mealybug; CENTER: Phenacoccus celtisifoliae (Hollinger), celtis mealybug; RIGHT: Nipaecoccus aurilanatus (Maskell), golden mealybug. Middle row. LEFT: Cataenococcus olivaceous (Cockerell), olive brown mealybug; CENTER: Rhizoecus pritchardi McKenzie, Pritchard mealybug; RIGHT: Spilococcus implicates Ferris, implicated mealybug. Bottom row. LEFT: Pseudococcus fragilis Brain, citrophilus mealybug; CENTER: Rhizoecus kondonis Kuwana, Kondo mealybug; RIGHT: damage by same to alfalfa.

COLOR PLATE IV. Top row. LEFT: Chnaurococcus trifolii (Forbes), clover-root mealybug; CENTER: Anisococcus quercus (Ehrhorn), interior live oak mealybug; RIGHT: Ferrisia virgata (Cockerell), striped mealybug. Middle row. LEFT: Scaptococcus milleri McKenzie, new species, Miller sand dune mealybug; CENTER: Pseudococcus maritimus (Ehrhorn), grape mealybug; RIGHT: Anisococcus oregonensis Ferris, nest mealybug. Bottom row. LEFT: Puto californicus McKenzie, new species, California puto mealybug; CENTER: Rhizoecus cacticans Hambleton, cacticans mealybugs; RIGHT: Puto acirculus McKenzie, grass puto mealybug (immature).

ECONOMIC erature. Until the paper by Wilkey and McKenzie (1961), no one knew exactly what species were being discussed. In California, P. maritimus is found primarily in the central and southern sections, and can be collected from the cool coastline to the hot central valleys. The presence of these mealybugs on the ripe marketed grapes results in serious economic loss to the growers. Heavy infestations cause the grapes to crack, allowing mold contamination (Ebeling, 1959). Present control of this mealybug on grapes is with parathion (Jensen, Stafford, and Break, 1959). Pseudococcus longispinus (Targioni Tozzetti), the long-tailed mealybug (col. pi. X V ) , is distributed world wide and is particularly common in America, Africa, and some parts of Europe (Ferris, 1950). This mealybug is often a pest in greenhouses and nurseries, but is also found out-of-doors in warmer areas. In California, the species is most important as a nursery pest, attacking a wide variety of hosts. It has occasionally been reported as a pest of avocados, grapes, and citrus. Specific plant damage is of the usual type. In the fall of 1933, a serious infestation of this mealybug developed on oranges over an area of some 200 acres near Downey in Los Angeles County, California. Although severe outbreaks are quite unusual for this species, some biological control work was done at that time, and partial control has been maintained since with the hymenopterous parasites, Anarhopus sydneyensis Timberlake and Tetracnemus peregrinus Compere (Flanders, 1940). Chemical control is feasible, especially in nurseries. Phenacoccus gossypii Townsend and Cockerell, the Mexican mealybug (col. pi. I ) , is widely distributed throughout the United States and Mexico. In California it is a pest of numerous flowering plants in nurseries and greenhouses, and in natural environments. This mealybug, which is most often found on the foliage of its host, apparently causes as much damage to its host as the citrus mealybug, Pianococcus citri (Risso) (Meyers, 1932). Wilting and stunting of the plant host are common effects of heavy infestations. Chemical control is feasible. Pseudococcus microcirculus McKenzie, orchid mealybug (col. pi. X V I I ) , is a species described rather recently (McKenzie, 1960), but one which has caused many problems to orchid growers before and since that time. It is restricted, as now known, to nurseries where it attacks only orchids. It is found primarily on the roots of its host, but crawls to the foliage and leaf sheaths when infestations become heavy. It is now restricted to California, ranging from the most serious infestations in the Marin County region south to Los Angeles and Riverside counties. Although most infestations have occurred in coastal regions, the controlled environments of the nurseries make even the hot valley regions susceptible. Restriction to California is probably more an indication of heavier collecting in this state than in others, for this mealybug was intercepted on orchids from Florida in 1961. The type of damage is the normal form of unsightly contamination with the pro-

IMPORTANCE

7

duction of honeydew during heavy infestations. (See col. pi. X V I I ) . Chemical control is partially successful, but because infestations are dften hidden in the sheaths and on the roots, the mealybugs avoid contact with the chemical. Chorizococcus brevicruris McKenzie, the shortlegged mealybug (col. pi. V I I I ) , was originally described (1960) from a single specimen intercepted in quarantine at Honolulu, Hawaii, on a succulent, Caralluma nebrowni (Asclepiadaceae), from an unknown California source. In September, 1965, this mealybug was found in a private greenhouse in Sacramento, California where it was causing serious damage to several species of succulents, Huerina spp. and Stapelia spp., belonging to the same plant family as the one intercepted in quarantine at Honolulu (see col. pi. VIII). It was also found on one cactus plant, Myrtillocactus geometrizans, growing near infested succulents. Recent survey indicates that this species is rather widespread in California nurseries in the counties of Los Angeles, Riverside, Sacramento, and San Bernardino. In 1941, it was collected on Stapelia sp. in a nursery in west Los Angeles, California, but was misidentified at that time as another pseudococcid species. The Sacramento grower reports that repeated applications of malathion, used as a drench for potted plants, is a fairly satisfactory control for this mealybug. Ferrisia virgata (Cockerell), the striped mealybug (col. pis. IV and X ) , has recently been of some concern to California economic entomologists. Until recently it was not known to occur in Califronia except in minor nursery infestations which were soon eliminated. Within the past few years, however, heavy infestations were noticed on many ornamental plants in Imperial County. This has caused some alarm, because in other parts of the world this species is reported as an important pest, especially to cotton. It is found normally above ground on the foliage where it causes the usual honeydew-sootymold-type damage. During severe weather, in Africa at least, it may move to the crown and roots of its host (De Lotto, 1964). In Imperial County a survey of the extent of infestation is presently being conducted by the California Department of Agriculture, Bureau of Entomology. Surveys have shown that the mealybug is found on a wide range of hosts with an apparent preference for lantana, Lantana spp. (Verbenaceae), "popper" plant, Ruellia malacosperma (Acanthaceae), blood flower, Asclepias curassavica (Asclepiadaceae), Chrysanthemum sp. (Compositae), and that the infestation is fairly localized. Investigations are being conducted to determine the advisability of eradication. Antonina graminis (Maskell), Rhodesgrass mealybug (col. pi. V I I ) , is another mealybug presently of major concern to economic entomologists in California. This mealybug has been a major pest of many grass species. Rhodesgrass, Johnson grass, Bermuda grass, and St. Augustine grass apparently are preferred hosts, but some 69 grass species have been recorded as hosts in the United States (Chada

8

ECONOMIC

IMPORTANCE

and Wood, 1960). It is a very serious pest on Rhodesgrass in the southwestern United States, where it has completely destroyed thousands of acres of good pasture land. Rhodesgrass mealybug is known to infest not only grassland areas but also residential lawns and golf greens. For the most part, this species is confined to the roots and crowns of its host, but attack of the aerial nodes does occur during heavy infestations. It is difficult to determine the extent of damage caused to the host because of the secondary effects of drought, overgrazing, or mowing. Injury is first indicated by the stunting and reduction in the overall size of individual grass clumps, with darkening of the leaves and eventual death of the host plant. Death of seedling plants is known to occur in about three weeks. Because of the serious threat that this mealybug poses to California lawns, pastures, and golf courses, entomologists have been carefully observing its behavior and spread in the state. The mealybug was first collected in Imperial County on October 13, 1957 on St. Augustine grass. On August 10, 1964 it was found in a lawn of Bermuda grass on a residential property at San Diego, California—a distance of some 85 miles from nearest infestation in Imperial County. Control is difficult because of the position of the mealybug on its host, and the use of the systemic insecticides Schradan and Demeton as a drench have proved effective only in certain confined areas. It is too expensive and impractical, however, to use such methods for large acreage infestations (Chada and Woods, 1960). Phenacoccus graminosus McKenzie, the ryegrass mealybug (col. pis. Ill and XII), is a species located primarily in the central and northern parts of California, ranging from the coast inland to the valleys. It is restricted to grass hosts where it primarily inhabits the grass sheaths. In California this mealybug is only of minor economic importance, but according to Miss Helen M. Brookes, Coccidologist at the University of Adelaide, Waite Agricultural Research Institute, Adelaide, South Australia, the species is of some importance on barley, canarygrass, ryegrass, and wheat in that country. In one instance the mealybugs were living in large colonies among the spikelets of Phalaris tuberosa (bulb canarygrass), feeding on the peduncle rather than on the ovaries. In this case they seem to have been responsible for the failure of many seeds to germinate. In New Zealand its habit of crawling from ground cover onto apple trees, settling around the stem end of fruits, has caused rejection of apple shipments to other countries. No control measures have been directed toward this mealybug. Trionymus haancheni McKenzie, the Haanchen barley mealybug, was reported by Osborn (1951), and later by McKenzie (1960), as of some economic concern to Haanchen barley grown in the Tulelake area of Siskiyou County, California. Osborn (1951, p.154) reports that in August of that year, "the Siskiyou County Agricultural Commissioner's office at Tulelake submitted specimens of a

mealybug reported to be so abundant on Haanchen barley in some fields that combines were being gummed up and clogged at harvest. When visited early in September, harvesting had been completed in the infested fields but large numbers of dead and dried mealybugs were present on the stubble. Large numbers of cottony egg clusters contained unhatched eggs. Broken straws on the ground and some standing uncut at the edge of the field showed evidence of feeding by mealybugs along the stems under the leaf sheaths. Accumulations of thick sticky honeydew secretions were still present. Approximately 15,000 acres of Haanchen barley are planted in this district, located south and west of Tulelake. Gumming of combines at harvest was said to have involved approximately 350 acres." Experimental control has not been directed against this mealybug. Heterococcus pulverarius (Newstead) (=H.graminicola Morrison), the bluegrass mealybug (col. pis. I and XI), has on several occasions become economically important on bluegrass grown for seed in northern California and southern Oregon. The mealybugs are found primarily in the grass blade sheaths, but when infestations become severe they may be exposed on the foliage. Honeydew produced by this species often causes the grass foliage to become quite sticky. When this happens at harvest time, it has been found necessary to first wet the windrows with water spray the day before combining. Chemical control, especially with parathion, has been used to control this mealybug. Cultural practices are also quite effective in holding the species in check. On certain occasions mealybugs may become rather troublesome, if not serious, household pests. During September, 1963, the fluffy mealybug, Puto echinatus McKenzie, became so abundant on vegetation growing in a residential area at Imperial Beach, San Diego County, California, that large numbers of specimens invaded dwellings much to the terror and dismay of the occupants. The preferred host of this coastal mealybug appears to be Australian saltbush, A triplex semibaccata (Chenopodiaceae), although during epidemic periods it may attack many other plant species. A similar situation developed during the summer of 1966, when the white mealybug, Puto albicans McKenzie, reached epidemic proportions on manzanita, Arctostaphylos sp. (Ericaceae), in an area situated approximately six miles southeast of Placerville, El Dorado County, California. As a result of this tremendous population explosion, great numbers of female mealybugs swarmed into homes in this area. Feeble attempts at control were made with no visible results, mainly because only a few plants were treated in an infested area of perhaps several square miles. Aside from numerous other species of mealybugs feeding above ground and causing economic concern to commercial, subtropical, and ornamental plants, there appears to be an enormous fauna of subterranean species about which very little is known.

ECONOMIC Hambleton (1946), in his studies of hypogeic mealybugs, records that two workers, Bunzli (1935) and Weber (1944), have demonstrated the importance of soil-inhabiting mealybugs in the production of certain tropical crops. No less than 10 species of ground mealybugs are associated with ants on roots of Coffea liberica in Dutch Guiana. Apparently, coffee and cacao are infested \yith so many species of subterranean mealybugs that it would be reasonable to assume that these insects will be found to play a more important role as pests than has previously been suspected. In California there are three species of groundinhabiting mealybugs of undertermined economic importance belonging to the genus Rhizoecus Klinckel d'Herculais. These species are: Rhizoecus falcifer KUnckel d'Herculais, the ground mealybug (col. pi. XXI); R. kondonis Kuwana, Kondo mealybug (col. pi. I l l ) ; and R. pritchardi McKenzie, Pritchard mealybug (col. pis. Ill and XXII). As mentioned in the chapter on Control Measures, the field species are almost impossible to control, whereas in gardens and nurseries flooding with water, or drenching potted plots in certain insecticides, has given fairly satisfactory control. Collections of Rhizoecus kondonis Kuwana, Kondo mealybug (col. pi. I l l ) , on roots of alfalfa, strawberry, and in leaf mold with uncertain plant associations, indicate that this species is more widespread in California than previously supposed, and that it is capable of causing serious economic damage to some of these commercial crops. Collections of Rhizoecus kondonis Kuwana were made during November, 1958, on the roots of alfalfa grown on a ranch eight miles west of Santa Rosa, Sonoma County, California. This mealybug was found aggressively feeding on alfala roots and causing extensive brown areas on killed and stunted plants. By June, 1959, approximately 50 per cent of the stand of alfalfa had been killed by the feeding activities of this mealybug. The land on which the alfalfa was grown was originally planted to Gravenstein apple. In 1929 the orchard was removed, and from that time until the fall of 1950 it was alternately sown to Sudangrass and oats. In 1951 the land was put into alfalfa and left in that crop for three years. Finally the alfalfa was removed, and each year thereafter, until the fall of 1956, the land was sown to oats. In the spring of 1957 it was again put into alfalfa. The alfalfa was grown on alluvial-type soil near a creek bed. It was nonirrigated, but subterranean water provided enough moisture to promote sufficient plant growth to justify three or four cuttings annually. The mealybugs were found to be feeding in great numbers just beneath the soil surface and to depths of more than 38 inches (col. pi. III). Small soil samples collected at various depths—8 inches, 12 inches, 24 inches, and 36 inches—were put into plastic bags and brought into the Entomology Insectary laboratory. These soil samples were then placed in four separate Berlese funnel traps. The number of

IMPORTANCE

9

mealybugs (nymphs and adults) recovered from the traps is summarized as follows: 8 inches = 3 specimens; 12 inches = 3 specimens; 24 inches = 12 specimens; and 36 inches = 23 specimens. No attempt was made to determine the number of mealybugs per cubic foot of soil in the alfalfa field. It is suspected that alfalfa growing in areas where frequent summer and fall irrigations are practiced may not be so susceptible to attack by this mealybug. Specimens of this same subterranean mealybug, Rhizoecus kondonis, collected from soil tube cores down to a depth of six feet beneath prune trees growing in the Yuba City area, Sutter County, California, were brought into the laboratory for identification by Dr. K. Uriu of the University of California, Department of Pomology, Davis. According to Dr. Uriu, the orchard from which the mealybugs were taken displayed zinc and potassium deficiency symptoms and, therefore, zinc and potassium plots were established to correct the situation. Apparently, the zinc symptoms were corrected, but the presence of yellow and scorched leaves typical of potassium deficiency was not altered. It is suspected that root injury produced by this mealybug could very likely be the factor in decline of these trees. Preliminary survey indicates that approximately 50 acres of prunes exhibited damage as described above. This problem of mealybugs on prune roots was brought to the attention of Dr. S. F. Bailey, of the University of California, Department of Entomology, Davis, who is responsible for research on insects attacking deciduous fruits. Dr. Bailey and I visited the affected prune acreage and took hand-dug soil samples to the three-foot level and soil tube core samples to the six-foot level to ascertain the mealybug population per cubic foot of soil. The infested soil samples were then subjected to Berlese funnel trap treatment to recover the mealybugs. The number of mealybugs (all stages) found per cubic foot of soil at various depths in these samples is recorded as follows: one foot =1,708; two feet = 1,245; and three feet = 870. The soil tube core samples taken at depths of four, five and six feet were not broken up before being put into the Berlese funnel trap; this resulted in the soil hardening and trapping the mealybugs before they could emerge. Mealybugs were, however, observed in all these soil samples before they were dried out. Hambleton (1946) recorded subterranean mealybugs commonly found associated with ants, and in their galleries accompanying grass roots, at depths of eight to ten inches below the surface. The information recorded above indicates their ability to feed effectively at greater soil depths than heretofore reported. Rhizoecus kondonis also appears to be a pest problem in commercial strawberry plantings. Attacked plants show marked stunting, foliage fading, and devitalization resulting in a commercially nonproductive planting. Rhizoecus pritchardi McKenzie, Pritchard mealybug (col. pis. Ill and XXII), has become a rather serious pest, especially to African violet grown under

ECONOMIC

10

IMPORTANCE

greenhouse and nursery conditions. Damage caused by this mealybug to African violet is manifested in devitalization of the plant, resulting in discoloration of the leaves, marked curtailment of blooms until no flowers are produced, and, in cases of heavy infestations, ultimate death of the plant itself (see col. pi. X X I I ) . This mealybug is rapidly spreading across the United States and has been reported as seriously damaging African violet in Maryland and Pennsylvania. Drenching the plants with certain insecticides has been employed as a method of control. Rhizoecus falcifer Kiinckel d'Herculais, the ground mealybug (col. pi. X X I ) , is a common subterranean species present in gardens, nurseries, and greenhouses throughout most of California. Damage caused by this species is general devitalization to complete death of hosts attacked. The mealybug seems to have no particular host preference. Partial control can be initiated by heavy flooding of the infested areas. In addition to these main problems in California, certain others are known in various parts of the world. Mealybug toxins are rather important in some areas. The pineapple wilt in the Hawaiian Islands involves the pineapple mealybug, Dysmicoccus brevipes (Cockerell). It is a serious economic problem to pineapple in that state and has resulted in an extensive amount of research in the past few years. Some virus diseases are also transmitted by mealybugs. Perhaps one of the most important of these diseases is swollen shoot of cacao. This disease is transmitted by several mealybug vectors, including Planococcoides njalensis (Laing), and Ferrisia virgata (Cockerell). This virus causes excessive damage to cacao trees each year. Mention is made by Newton ( 1 9 5 3 ) of transmission of tobacco mosaic virus by Planococcus citri (Risso). Also it is recently believed that the pineapple wilt mentioned above as a toxin is in some way connected with a virus. FIELD KEY TO COMMON CALIFORNIA MEALYBUGS OF ECONOMIC IMPORTANCE 1.

2(1).

3(2). 4(3).

Wax filaments present around body margin 2 Wax filaments absent or restricted to two or three caudal pairs 8 Dorsum with at least one longitudinal stripe 3 Dorsum of uniform color, without stripe 6 With a single dorso-medial stripe 4 With two dorsal sublateral stripes . . . . 5 Caudal pair of wax filaments elongate, normally as long as or longer than total length of body Pseudococcus longispinus (Targioni Tozzetti) (see col. pi. XVI) (long-tailed mealybug) Caudal pair of wax filaments approximately same length as others, at most one-quarter of total length of body . . Planococcus citri (Risso) (see col. pis. I and X I V ) (citrus mealybug)

5(3).

6(2).

7(6).

8(1).

9(8).

10(9).

Body contents when crushed bright red; lateral wax filaments easily visible . . Pseudococcus fragilis Brain (see col. pis. I l l and X V ) (citrophilus mealybug) Body contents when crushed gray to light pink; lateral wax filaments minute, difficult to see Phenacoccus gossypii Townsend & Cockerell (see col. pi. I ) (Mexican mealybug) Wax filaments small; found on grass . . . . . . Phenacoccus graminosus McKenzie (see col. pis. I l l and X I I ) (ryegrass mealybug) Wax filaments elongate; not found on grass 7 Wax filaments wavy; always found on orchids . . Pseudococcus microcirculus McKenzie (see col. pi. X V I I ) (orchid mealybug) Wax filaments straight; rarely if ever found on orchids . . . Pseudococcus maritimus (Ehrhorn) (see col. pi. I V ) (grape mealybug) Body size small; soil inhabiting; antennae elbowed Rhizoecus spp. (see col. pis. Ill, X X I and X X I I ) (rhizoecus mealybugs) Body size normal; foliage inhabiting; antennae straight 9 Dorsum with two longitudinal stripes and many long crystalline rods . . . . Ferrisia virgata (Cockerell) (see col. pis. IV and X ) (striped mealybug) Not as above 10 Legs absent; enclosed in a white felted sac; with a long white tubular filament usually protruding from anal opening in sac Antonina graminis (Maskell) (see col. pi. V I I ) (Rhodesgrass mealybug) Not as above. [This half of the couplet involves two species which are very similar in appearance. They both are found in grass blade sheaths. Trionymus haacheni McKenzie, (Haanchen barley mealybug) is normally found on Hordeum vulgarae, while Heterococcus pulverarius (Newstead) (bluegrass mealybug) is rarely if ever found on the above host (see col. pis. I and XI).] REFERENCES

Basinger, A. J. 1931. Field key for the determination of some of the common mealybugs infesting nursery stock in California. Calif. Dept. Agr. Bull. 20(2) :189—193, illus.

ECONOMIC Bunzli, G. H. 1958. Untersuchungen über coccidophile Ameisen aus den Kaffeefeldern von Susiman mitt. Schweiz. Ent. Ges. 1 6 ( 6 - 7 ) :455-593. Chada, H. L. and E. A. Wood. 1960. Biology and control of the Rhodesgrass Scale. U. S. Dept. Agr. Tech. Bull. 1221, 21 pp., illus. Compere, H. and H. S. Smith. 1932. The control of the citrophilus mealybug, Pseudococcus gahani, by Australian parasites. Hilgardia 6 ( 1 7 ) : 585-618. DeBach, P. and E. I. Schlinger. 1964. Biological control of insect pests and weeds. Chapman and Hall Ltd., London. 844 pp., illus. De Lotto, G. 1964. Observations on African Mealybugs (Hemiptera: Coccoidea). Bull. Brit. Mus. (Nat. Hist.) Ent. 1 4 ( 8 1 ) : 3 4 3 - 3 9 7 , illus. Ebeling, W. 1959. Subtropical fruit pests. Univ. of Calif. Div. Agr. Sei. Calif. 436 pp., illus. Ferris, G. F. 1950. Atlas of the scale insects of North America (series V). The Pseudococcidae (Part I) Stanford Univ. Press., Stanford. 278 pp., illus. Flanders, S. E. 1940. Biological control of the longtailed mealybug, Pseudococcus longispinus. Jour. Econ. Ent. 33(5):754-759. Hambleton, E. J. 1946. Studies of Hypogeic Mealybugs. Rev. de Ent. 1 7 ( 1 - 2 ) : 1-77, illus. Jensen, F., E. M. Stafford, and R. A. Break. 1954. T h e Grape Mealybug. Calif. Agr. 8 ( 3 ) : 3 3 0 .

IMPORTANCE

11

McKenzie, H. L. 1960. Taxonomic study of California mealybugs with descriptions of new species. (Homoptera: Coccoidea: Pseudococcidae). Hilgardia 2 9 ( 1 5 ) : 681-770, illus. Myers, L. E. 1932. Two economic greenhouse mealybugs of Mississippi. The citrus mealybug and the Mexican mealybug. Jour. Econ. Ent. 2 5 ( 4 ) : 8 9 1 896, illus. Newton, W. 1953. Transmission of tobacco mosaic by citrus mealybug. F.A.O. Plant Prot. Bull. 2 ( 3 ) :40. Osborn, H. T. 1951. Insect pest survey. Thirty-second annual report, period ending December 31, 1951. Calif. Dept. Agr. Bull. 4 0 ( 4 ) : 154. Pritchard, A. E. 1949. California greenhouse pests and their control. Calif. Agr. Expt. Sta. Bull. 713. 71 pp., illus. Quayle, H. J. 1938. Insects of citrus and other subtropical fruits. Comstock Publishing Co., Ithaca. 583 pp., illus. Weber, N. A. 1944. The neo-tropical coccid-tending ants of the genus Acropyga Roger. Ann. Ent. Soc. Amer. 37(1):89-122. Wilkey, R. F. and H. L. McKenzie. 1961. Systematic status of the Pseudococcus martimus-malacearum complex of mealybugs (Homoptera: Coccoidea: Pseudococcidae). Calif. Dept. Agr. Bull. 5 0 ( 4 ) : 245-249, illus. Woglum, R. S. 1922. Control of the citrophilus mealybug. U.S.D.A. Farmer's Bull. 1040. 4 pp.

CONTROL MEASURES

CONTROL

Mealybugs throughout the world cause a variety of economic problems. The most obvious damage is caused by the sucking habits of these insects. Heavy infestations often cause stunting or death to the plant host. At times mealybugs have toxins and act as vectors of certain viruses detrimental to plant life (Newton, 1953). The young and adults excrete quantities of honeydew which attracts ants and serves as a medium of growth for sooty mold, causing serious smutting of the fruit and foliage. These types of plant contamination are not only important to such crops as alfalfa, citrus, grapes, pears, and avocados, but they are also detrimental to ornamental host plants. Mealybugs are a major problem in California greenhouses and nurseries, where they often cause severe economic damage. The frequent use of insecticides and labor for mealybug control has made their cost to the grower still greater. It seems surprising that such a delicate soft-bodied insect should be so difficult to control. There are, however, several reasons which may account for this fact. Perhaps the most important factor is the habitat of the mealybug. Mealybugs live in protected areas such as bark cracks and crevices, in grass sheaths, at bases of leaf petioles, and on the undersides of leaves. In addition, other species have the habit of spending their entire lives deep in the soil, protected almost entirely from insecticidal materials. Also the eggs of mealybugs, protected by the waxy filamentous secretions of the ovisac, are almost impossible to reach with insecticides. As a consequence, this is perhaps one of the least susceptible stages of mealybug development to effective control, except for some insecticides that act as both contact and fumigant poisons, i.e., parathion and dichlorovos. Some mealybugs have been able to develop resistance, while still others are just not affected by certain insecticides.

It is not my purpose to give specific recommendations for mealybug control, but rather to briefly summarize the more recent control measures, resulting from experimentation by other workers, some of which have proven quite effective. Therefore, a résumé of mealybug control measures as gleaned through a search of the literature is here presented. Insecticide trade names mentioned in the following paragraphs are given so that the material may prove more intelligible to the reader. No discrimination is implied with respect to similar or other available preparations. Organic

Phosphates

No one specific insecticide is effective against all species of mealybugs, but parathion comes the closest to fulfilling this aim (Pritchard, 1949). It should be noted, however, that parathion is used primarily in commercial agriculture and is not available to backyard users. Parathion is a contact and stomach poison having some fumigant effect and is used as both a dust and a spray. Madsen (1962) pointed out that, at least for the control of Pseudococcus maritimus (Ehrhorn), dusts were much less effective than sprays. Although this insecticide has been successful in mealybug control, it is extremely hazardous and should be handled with much care. Because of the hazard of parathion, and because some resistance has been noted in connection with this insecticide (Berry, 1961 MS.), a search was made for a replacement. Malathion, which is primarily a contact insecticide, has proved to be a good substitute for it. Although caution should be used in applying malathion, it is certainly much less dangerous than the first mentioned insecticide. Malathion is used primarily in the control of garden and nursery 12

CONTROL mealybugs (Michelbacher, 1959) and is effective in controlling several species, including Ferrisia virgata (Cockerell) (Highland, 1956), Heterococcus pulverarius (Newstead) (Dietz, 1960), Dysmicoccus obesus (Lobdell) (Milliron, 1958), Dysmicoccus brevipes (Cockerell) (Carter, 1952), and Dysmicoccus cuspidatae (Rau) (Weidhaas and Shaw, 1956). Apparently, Planococcus citri (Risso) is not effectively controlled with this insecticide (Kriegler, 1957). Diazinon is another insecticide which is effective at least in the control of Pseudococcus maritimus (Ehrhorn) (Madsen, 1956; Madsen and McNelly, 1960; and Madsen and Westgard, 1962). It is believed that this material is also effective against other mealybugs. Effective both by contact and stomach action, it is used as a dust and a spray, and is not as hazardous as parathion. TEPP, another organic phosphate, is primarily a contact-type insecticide. It is extremely hazardous and seems to be fairly ineffective in mealybug control when used as a spray (Richardson, 1953). When this compound is used as a smoke, however, it has proven quite successful in greenhouses against immature stages of mealybugs, particularly if a series of applications are made. This method has effectively controlled the following species of mealybugs: Planococcus citri (Risso), Phenacoccus gossypii Townsend and Cockerell, Pseudococcus longispinus (Targioni Tozzetti), and P. maritimus (Ehrhorn) (Jefferson and Mack, 1953; and Jefferson ard Pritchard, 1961). Dichlorovos (DDVP) is also registered for use in greenhouses as a contact and fumigant control of certain mealybug species. Several organic phosphates have been used primarily as systemic insecticides. Some of these which have proven effective in mealybug control, in limited cases, are phorate (Thimet®) 1 and DiSyston®. The former has been used in the control of Heterococcus pulverarius (Newstead) (Dietz and Harwood, 1960), Ferrisia virgata (Cockerell), Planococcoides njalensis (Laing) (Bowman and Casida, 1958), and Pseudococcus comstocki (Kuwana) (Abrahao and Manprim, 1958). The latter has been used in the control of Dysmicoccus brevipes (Cockerell) (Carter and Gortner, 1958). The application of these chemicals for control of the mealybugs mentioned above has been either as granules in the soil, as a spray on the soil, or by "injection" through borings in the plant. Both insecticides are highly hazardous. Dimefox is another systemic organic phosphate and has been used successfully in Africa to control Planococcoides njalensis (Laing). It is, however, phytotoxic to some plants (Bowman and Casida, 1958), and consequently its usage is limited. Other systemic organic phosphates which have been recently unsuccessful are systox and Schradan (OMPA). Both were used without success in control 1

® = registered trade names.

MEASURES

13

programs of Dysmicoccus boninsis (Kuwana) (Charpentier, 1956; Carter, 1952). Chlorinated Hydrocarbons DDT has been used extensively for the control of many insects, and mealybugs are included on this list. Within the past ten years DDT has been only slightly effective against mealybugs. In fact, it has been recorded as ineffective against Maconellicoccus hirsutus (Green) (Ankersmit, 1951), Pseudococcus maritimus (Ehrhorn) (Madsen, 1956), and Antonina graminis (Maskell) (Richardson, 1953); and in the case of Planococcus citri (Risso) an increase in the mealybug population may occur (Bartlett, 1957). Other chlorinated hydrocarbons which may occasionally have a moderate effect on the control of mealybugs are lindane, aldrin, dieldrin, and endrin. Lindane, as recently indicated in literature, has been effective in three mealybug control programs and ineffective in four. In one of the successful programs this compound was used as a fumigant against Pseudococcus maritimus (Ehrhorn) on potato sprouts being raised for seed. Aldrin seems most effective when used in combination with dieldrin or lindane with chlordane as an addition for ant control (Fernandez, 1957). Although some attempts have been made with aldrin alone, these tests were ineffective. Aldrin is used as a contact and stomach poison and may have some fumigant action. Dipping young plants in combination formulations and then spraying the plants at regular intervals with these formulations seem to be fairly successful (Abrahao and Andrade, 1961). Aldrin may occasionally be phytotoxic and is moderately hazardous. Dieldrin and endrin have been used in much the same manner as aldrin and give approximately the same amount of success. Both should be handled the same as aldrin. Chlordane is often used in combination with other insecticides, but when used alone has little effect in mealybug control (Richardson, 1953). Most often chlordane is used to kill ant attendants which keep parasites and predators from harming their mealybug hosts. Oftentimes effective control results from controlling the ants and allowing the natural parasites to control the mealybugs (LePelley, 1953). Another unsuccessful chlorinated hydrocarbon which has been used in mealybug control in the past is toxaphene (Baccolo, 1953). It proved ineffective against Pseudococcus spp. (Willie, Simon, and Gonzales, 1955), Planococcus citri (Risso) (Baccolo, 1953), and Maconellicoccus hirsutus (Green) (Ankersmit, 1951). Ground Mealybug Control The control of soil mealybugs remains to be solved. The problems with Rhizoecus pritchardi McKenzie and R. kondonis (Kuwana) are examples of the need of control of these pests. Some partially sue-

14

CONTROL

MEASURES

cessful controls have been found. Where soil is known to be infested, and if all plants in that soil are expendable, control has in the past been gained by spraying the newly spaded earth with a dichloroethyl ether solution (Michelbacher, 1959). This insecticide is rather old and may no longer be available. Malathion and parathion drenches are often used and are partially effective. It has been reported in correspondence that heavy infestations of Rhizoecus pritchardi McKenzie on roots of African violet have been successfully controlled by drenching the potted plants in either dimethoate (Cygon®) or VC-13®, giving them two applications at a two-week interval. Aldrin, used as a drench, also gave excellent control of this pest (Snetsinger, 1966). Miscellaneous Chemical Control Summer oil emulsions are particularly effective in the control of mealybugs on ornamental plants. Applications should be made at regular intervals of one to three weeks (Michelbacher, Swift, Davis, Hall, and Raabe, 1959). Combinations of these oil emulsions with contact insecticides are quite effective in the control of garden and household mealybugs. Insecticides which have proved virtually unsuccessful for mealybug control are: lead arsenate (Madsen, 1956), nicotine, rotenone (Ankersmit, 1951), HCN when used against Planococcus citri (Risso) (Constantino, 1956) and ryania (Patterson and MacLellan, 1955). Adversities Caused by Chemicals Although it is apparent that many problems arise when chemicals are applied for control of any insect, certain of these aftereffects warrant special mention in the case of mealybug control. Oftentimes when application of insecticides are made for the control of certain mealybugs, instead of causing a decrease in the pseudococcid population, an increase in noted. This is usually due to the fact that the insecticides have little effect on the mealybug, but eliminate the natural enemies which were at one time holding the pseudococcid population in check. At times certain insecticides will control the mealybugs, but, with the decrease of competition, other pests, particularly mites, will increase. This second pest may become a more serious pest than the first. Finally, it has often been noted that, although a chemical may give excellent control for a short time, the pseudococcid population when building up again will reach an even higher level than it had attained before chemicals were first applied. Cultural Methods Other types of control measures are effective in some cases. Crop sanitation is useful in control of Heterococcus pulverarius (Newstead) (Dietz, 1960), and H. nigeriensis Williams (Harris, 1961). Both of these insects are pests on grass where they inhabit the grass sheaths, making chemical contact

nearly impossible. Burning and plowing the crop under after harvest results in very little reoccurrence of the mealybugs. Michelbacher (1959) mentions that the control of certain garden mealybugs may be effected simply by hosing the plants down with a strong stream of water. Although this seems rather unorthodox, control is fairly successful, especially if this treatment is used at regular intervals. Biological control and combinations of biological and chemical controls are another very important area of mealybug suppression. In fact, mealybugs are one of the main subjects of biological control, and a tremendous amount of research has been done in this area, much of it quite successfully. The advantages and importance of biological control of pseudococcids cannot be stressed too much, but this is a field too complex to be treated in a work such as this. SUMMARY 1. Mealybugs are pests because they suck plant juices, have toxins, act as vectors of viruses, produce honeydew, and act as primary hosts for other pests. 2. Mealybugs are difficult to control because they inhabit protected areas, cover their eggs with a waxy ovisac, and are themselves covered with protective waxes. Some have developed resistance to certain insecticides. 3. Only a few chemicals are effective against mealybugs. Parathion seems to be the most effective but is extremely hazardous. In gardens, malathion or oil emulsion sprays or combinations of malathion and oils work quite well. Dichlorovos and diazinon are also useful for mealybug control. 4. Some systemic and fumigant insecticides are useful in certain specialized cases. 5. Biological control has proved to be fairly successful in mealybug control. 6. Crop sanitation and miscellaneous controls are sometimes successful. 7. Control of ground mealybugs is difficult. Applications of dichloroethyl ether, dimethoate, VC-13®, aldrin and malathion or parathion drenches have been partially successful. 8. When using chemicals be sure handling precautions are taken. Be aware of the aftereffects of chemical control. REFERENCES Abrahao, J. T. and O. Mamprim. 1958. Conchonilha da raiz do cafeeiro. Biologico 24(12):268-271. Abrahao, J. T. and O. C. Andrade. 1961. Decadencia do abacaxi causada pelo piolhobranco. Biologico 27(10):237-241. Ankersmit, G. W. 1951. Bestrijdingsproven tegen da katoenboorder Earias fabia Stoll. (tepidoptera, Noctuidae) Contr. Gen. Agr. Res. Sta. Bogor, no. 126. 19 pp.

CONTROL Baccolo, S. 1953. Experimente di lotta contro Pseudococcus citrì Risso. Biol. Zool. Agr. Bachic. 18(2— 3 ) : 107-115. Bartlett, B. R. 1957. Biotic factors in naturai control of citrus mealybugs in California. Jour. Econ. Ent. 50(6):753—755. Berry, D. W. 1961. Bionomics of the mealybug attacking pear trees in northern California. (Unpublished thesis for Masters Degree, Univ. of Calif., Davis.) Bowman, J. S. and J. E. Casida. 1958. Systemic insecticides for Theobroma cocoa L., their translocation and persistence in foliage and residues in cocoa beans. Jour. Econ. Ent. 51(6) :773-780. Carter, W. 1952. Organic phosphates as systemic insecticides on pineapple plants. Jour. Econ. Ent. 4 5 ( 6 ) : 981-984. Carter, W. and W. A. Gortner. 1958. The translocation of radioactive (S 35 ) Bayer 19639 in pineapple plants. Jour. Econ. Ent. 51(6):905-907. Charpentier, L. J. 1956. Systemic insecticide studies for control of vectors of sugar cane mosaic in Louisiana. Jour. Econ. Ent. 49(3) :413-414. Costantino, G. 1956. La lotta contro la cocciniglie degli agrumi mediante emulsioni di olii minerali e di esteri fosforice, da sole o in miscele. (Resultati di prove sperimentali.) Notiz. Mal. Piante 37-38 (16-17):91-123. Dietz, S. M. and R. F. Harwood. 1960. Host range and damage by the grass mealybug Heterococcus graminicola. Jour. Econ. Ent. 53(5) :737-740. Fernandez, A. 1957. Las "escamas palomillas" del cafeto en Venezuela y avances logrados en su control quimico. Agron. Trop. 7(1) :7—17. Harris, E. 1961. Distortion of guineacorn (Sorghum vulgare) caused by a mealybug Heterococcus nigeriensis Williams, in northern Nigeria. Bull. Ent. Res. 51(4) :667-684. Highland, H. A. 1956. The biology of Ferrisiana virgata, a pest of azaleas. Jour. Econ. Ent. 49(2) -.216-211. Jefferson, R. N. and G. E. Mack. 1953. Control of certain greenhouse insects and mites with tetraethyl dithiopyrophosphate smokes. Jour. Econ. Ent. 46(1) : 120-123.

MEASURES

15

Jefferson, R. N. and A. E. Pritchard. 1961. Pest control guide for California greenhouse plants and flowers. Calif. Agr. Expt. Sta. Ext. Ser. Leaf. 134. Jensen, F., D. Flaherty, E. M. Stafford, and H. Kido. 1964. Developments in control of the grape mealybug. Jour. Econ. Ent. 57(3):372-374. Kriegler, P. J. 1957. Malathion versus parathion for chemical control of the mealybug Planococcus citri (Risso) on vines. Jour. Ent. Soc. So. Afr. 2 0 ( 1 ) : 187188. Le Pelley, R. 1953. Insecticideal treatments against the mealybug ant (Pheidole punctulata) on coffee. Monthly Bull. Coffee Bd. Kenya 18(216) :569. Madsen, H. F. 1956. Codling moth control on pears, 1954-1955. Jour. Econ. Ent. 49(4) :467-470. Madsen, H. F. and L. B. McNelly. 1960. Control of the grape mealybug on apricots. Jour. Econ. Ent. 53(3) :354-357. Madsen, H. F. and P. H. Westgard. 1962. Behavior and control of the grape mealybug on pears. Jour. Econ. Ent. 55(6):849-850. Michelbacher, A. E., J. E. Swift, C. S. Davis, D. H. Hall, and R. D. Raabe. 1959. Ridding the garden of common pests. Calif. Agr. Expt. Sta. Ext. Serv. Circ. 479:30-31. Milliron, H. E. 1958. Economic importance and control of Loblolly mealybug Dysmicoccus obesus Lob. Jour. Econ. Ent. 51 (4):555-556. Newton, W. 1953. Transmission of tobacco mosaic by citrus mealybug. F. A. O. Plant Prot. Bull. 2(3) :40. Patterson, N. A. and C. R. MacLellan. 1955. Control of the codling moth and other orchard pests with ryania. 85th Rep. Ent. Soc. Ont., pp. 25-32. Pritchard, A. E. 1949. California greenhouse pests and their control. Calif. Agr. Expt. Sta. Bull. 713:21-23. Richardson, B. H. 1953. Insecticidal control of Rhodes grass scale on St. Augustine grass lawn. Jour. Econ. Ent. 46(3) :426-430. Snetsinger, R. 1966. Biology and control of a root-infesting mealybug on Saintpaulia. Jour. Econ. Ent. 5 9 ( 5 ) : 1077-1078. Weidhaas, J. A. and F. R. Shaw. 1956. Control of the Taxus mealybug with notes on its biology. Jour. Econ. Ent. 49(2):273-274. Willie, J. E., J. E. Simon, and J. E. Gonzales. 1955. Los insectos e insecticidas en la algodonera campana 1954-1955. Inf. Estac. Exp. Agr. La molina, no. 97. 20 pp.

ECOLOGY Mealybug Habitats, Ant Associations

MACRO-ENVIRONMENTAL PREFERENCES

The general groups of plants infested are perennial- and annual-flowering plants—(angiosperms), grasses, conifers, and some ferns. These along with inanimate objects where mealybugs are frequently found, and multiple host positions, are discussed on the following pages.

Mealybugs are much more numerous than might be expected. Most people know of them as common garden and nursery pests, but few realize the enormous native fauna which exists. In California mealybugs are found from the low moist coastal regions to the high snow-covered Sierra. They are found from elevations of -200 feet at Saltan Sea in Riverside County to altitudes of over 12,000 feet above Tuolumne Meadows in Mariposa County. They are found in salt marshes and hot, dry desert sands. In these different environments certain types of mealybugs are found more frequently than others. In the very high altitude regions, conifer- and perennial-inhabiting Puto and grass sheath-infesting Trionymus are most likely to be encountered. In the moist coastal regions, root- and soil-inhabiting Rhizoecus and grass-infested Trionymus are common. In the dry, arid deserts, foilage- and rootinhabiting Phenacoccus and Spilococcus are numerous. These regions are not restricted to the genera mentioned, nor are the genera restricted to just these regions, but if a person were to collect in the mentioned areas these mealybugs would be the ones most likely encountered. There are some mealybugs which have no particular environmental preference. Amonosterium lichtensioides (Cockerell) is a good example of this, for its only restriction is the distribution of its host, Artemisia (Compositae) (see col. pi. VI). It is found in high mountains at altitudes of 10,000 feet to low coastal regions of 30 feet. It also is commonly found in dry chaparral areas.

Perennial-Flowering Plants (Excluding Grasses) This group is one upon which mealybugs are most commonly collected. In some regions every possible niche is utilized on the perennial hosts. In one instance in San Diego County, an Artemisia plant was infested with Rhizoecus gracilis McKenzie on the roots, Phenacoccus antemisiae Ehrhorn on the crown, Spilococcus corticosus McKenzie, herein described as new, under the bark, and Amonostherium lichtensioides (Cockerell) on the foliage. It is rare when so many positions on a plant are inhabited, but this case illustrates the possible areas of a plant which may be infested with pseudococcids. Perennial foliage-infesting mealybugs, although common in the field, are predominately noticed in greenhouses and backyard gardens. On foliage and stems mealybugs apparently prefer tight enclosed areas, for they normally are found in leaf or stem axils. Infestations on fruit are almost always enclosed in the calyx area. On the foliage such mealybugs as Planococcus citri (Risso), Pseudococcus longispinus (Targioni Tozzetti), and P. obscurus Essig are common (see col. pis. XIV, XVI, and XVIII). Stems and twigs of perennials are also good areas for mealybug colonies. Normally the mealybugs are found wedged into the cracks and crevices of the bark, again showing a preference for tight areas. Stems and twigs of plants are not nearly as important as roots, crowns, and leaf axils, but such mealybugs as Spilococcus eriogoni (Ehrhorn), Phenacoccus defectus Ferris, and P. alleni McKenzie are occasionally found restricted to the branches. Most

MICRO-ENVIRONMENTAL PREFERENCES Not only do mealybugs show certain macro-environmental choices, but the wide diversity of host plants and plant positions, or micro-environmental preferences (col. pi. V), are also important. 16

COLOR PLATE V. MEALYBUG HABITATS: leaves (including grass sheaths), stems, bark, crowns, roots, under rocks, and in ant galleries.

ECOLOGY

often branches are an overflow area from heavily infested leaf axils. Crown-infesting mealybugs are very numerous and are perhaps the most common of our native pseudococcid fauna. In this context the crown region is considdred to be the area of infestation on the plant which is from two to three inches above, to about the same distance below, the soil surface. Because of this, .the crown region may be divided into two parts: that part which is above ground and that which is below. The most important of these two groups is the subterranean crown. In this region very heavy infestations often occur (see col. pis. XIII and XXIII). Frequently, this area becomes so heavily contaminated with mealybugs that an over flow often occurs onto the roots. It should be noted that only very rarely does this overflow move to the crown above the soil surface. Infestations of moderate- to smallsized mealybugs usually occur in cracks and crevices on the crowns, whereas large species such as Puto yuccae (Coquillett) or Puto decorosus McKenzie, herein described as new, usually just cling to the bark in any way possible. Mealybugs of the Phenacoccus type may also be found encysted in the center of the trunk of the perennial plant itself. The only way that these mealybugs can be collected is by pulling the crown apart, .piece by piece. Perhaps these pseudococcids gain entrance into this area through cracks in the bark, but they appear to be completely enclosed by the plant tissue. Phenacoccus artemisiae Ehrhorn is an example of this type of mealybug. Very often the crown both above and below ground is utilized by certain boring insect larvae. Commonly if these boring tunnels are pulled open, heavy infestations of certain species of mealybugs will be discovered. Phenacoccus eremicus Ferris seems to depend upon the cerambycid boring tunnels for its total existence during detrimental weather. When snow is on the ground or temperatures are consistently above 100°F., these mealybugs are found only in the cerambycid tunnels. Apparently the tunnels are an ideal situation for this pseudococcid, for the empty cavities surrounding the beetle larva are quite often completely full of mealybugs in all stages of development. During more favorable weather, infestations outside of the tunnels are quite common. Root infestations are also common, especially on our native flora (see col. pi. V). Many species of mealybugs are entirely restricted to the root region. These mealybugs can be divided into two types: those which inhabit the main, large roots; and those which restrict themselves to the small fleshy roots. Cracks and crevices are the common areas of infestation on the main roots, but on the fleshy roots infestations may occur in any area where there is a vacant space. Species commonly collected in the large root areas are Phenacoccus solani Ferris, P. artemisiae Ehrhorn, Chorizococcus polyporus McKenzie, Puto pacificus McKenzie, herein described

17

as new, Chnaurococcus trifolii (Forbes), and many more. Species found in the second group are usually representatives of the genus Rhizoecus, with R. falcifer Kiinckel d'Herculais (see col. pi. XXI), R. gracilis McKenzie and R. kondonis Kuwana as good examples. Annual-Flowering

Plants (Grasses

Excluded)

Mealybugs infesting annual plants are unique in that their host is present for only a small part of the year. There is a question about the location of these mealybugs during that part of the year when their host plant is absent. This is still a mystery, although some casual field investigations have been made pertaining to the subject. In one instance a very heavy infestation of Phenacoccus eschscholtziae McKenzie was collected in early April on a large variety of annual hosts. Two months later an attempt was made to collect the mealybug again at exactly the same locality. All plants in the area were carefully examined, but no specimens were found. Soil samples were taken at depths of approximately three feet under hosts known to have been previously infested, and this soil was later placed in a Berlese funnel. No signs of mealybugs were found. It seems quite likely that the eggs remained on the dried host until new annuals emerged the next year. There is, however, no evidence to support this assumption. When the dried host plants, which obviously had been heavily infested a few months previously, were examined, only a very few, probably inviable, eggs could be found; the rest had apparently hatched. So for Phenacoccus eschscholtizae, at least, and probably other annual-infesting mealybugs, the disappearance of the pseudococcid with its host, is still a mystery. Infestations on annual hosts are normally localized in the subterranean areas, but there are mealybugs that are commonly found on the foliage of such hosts. Spilococcus atriplicis (Cockerell) would be an example of this. Grasses

Grasses are excellent hosts for many mealybugs (see col. pis. V, XI, XII, XXI, and XXV). They harbor not only large numbers of species but also numerous odd genera. Mealybugs are found on grasses in three areas: the foliage, crowns, and roots. Foliage infestations differ from those previously mentioned in that most of the mealybugs inhabit the leaf-blade sheath of the grass plant. Mealybugs inhabiting this area are not often seen, for they are completely hidden in the grass sheath and may be exposed only by removing the sheath from the plant. This type of pseudococcid is especially adapted to its habitat in that it is dorso-ventrally flattened, allowing it to fit comfortably in its sheath environment. Trionymus is the most common genus of this type with T. dolus Ferris, T. smithii (Essig), and T. festucae (Kuwana) being good examples. There are several grass-infesting species which are recorded as being exposed on the leaf surfaces of

18

ECOLOGY

the Gramineae. Again Trionymus is a good example of this type of infestation, but at times Heterococcus arenae Ferris, Distichlicoccus salinus (Cockerell), Chorizococcus rostellum (Hoke), and Phenacoccus graminosus McKenzie (see col. pi. XII), may be found exposed on the leaves. Many species of Discococcus are commonly found in the crown regions of Gramineae. This genus is probably quite common, but since it blends so well with its grass hosts, it is often overlooked. The adult females, which are fairly large and rotund, are found in small niches formed between the sheathing-leaf bundles in perennial grasses. They are also difficult to see because their coloring, which ranges from orange to light green, is usually similar to that of the grass host. Important also are the Antonina mealybugs, which are commonly found at the nodes of the grass stems. Antonina graminis (Maskell), an important pest of Rhodesgrass, is particularly well known for its habit of collecting around the aerial nodes of the plant (see col. pi. VII). Many types of mealybugs are found on the roots. Such odd genera as Misericoccus, Syrmococcus, and Cryptoripersia should be mentioned along with the more common Rhizoecus type genera. Most of these mealybugs feed on the tender terminal roots of the grass, where they are exposed to the surrounding soil with little or no waxy protection. Cryptoripersia, on the other hand, occurs in the same area on the roots, but is covered by a tough, felted sac (see col. pi. IX). Another root area is inhabited by Discococcus spectabilis McKenzie. The mature adult females of this mealybug form their ovisac at the base of the plant just a few millimeters below where the roots were first produced from the culm base. Conifers

The conifer-infesting mealybug habitats are not too well known, and practically no information is available pertaining to their full life cycle. The genus Puto contains several species which inhabit various types of conifers. What is surprising about these mealybugs is the fact that most of them have been found only in cracks in the bark or under rocks and fallen logs (see col. pi. V). This host position is quite extraordinary because it would seem virtually impossible for the mealybugs to gain any nourishment from such inanimate objects. Some of these pseudococcids have been found on the bark at the base of the trunks of redwood trees well over 150 feet tall. The bark in this area would undoubtedly be over ten times thicker than the length of the mealybug stylets. Considering the fact that gaining nutrition from bark is virtually impossible, some attempt was made to locate the areas where nutrient substances might be taken in by the mealybugs. All hosts in the vicinity were carefully examined, conifer roots and foliage

were checked, and soil samples were taken; none of these areas gave results. In light of this it was felt that perhaps these later instar nymphs were able to sustain themselves completely on nutrients gathered in the early parts of their life history. Perhaps the roots or the foliage of the conifer is the area of early development, and the bark is merely a resting place for later development where no feeding occurs. With this possibility in mind, about ten specimens of Puto laticribellum McKenzie were collected on the bark of Pinus ponderosa on July 13, 1964. These specimens, all later instar nymphs, were placed in a small cardboard box where they remained without any kind of nutrients. By November 15, the mealybugs had shed their skins and developed into mature male and female pseudococcids. It is, however, important to realize that the female specimens were quite reduced in size compared with normal P. laticribellum. Because of their abnormal size it is concluded that nutrients assimilated during the earlier nymphal stages are not solely adequate for complete development of the mealybug. Washburn (1965) shed some light on this subject in his work on Puto sandini Washburn which infests Englemann spruce, Picea engelmanii, in Utah, at elevations of 10,000-11,200 feet. He found that this mealybug goes through a very complicated four-year life cycle involving annual multiple migrations. As a demonstration of the complexity of the life cycle, here is what happens during the first year. The mature adult females produce the first instar nymphs under bark chips on the bole of the tree. These nymphs migrate from the bole to the duff at the base of the tree in late September where they remain under the snow until sometime in May. At this time they migrate back up the tree to the foliage where they feed until mid-July. At this time they migrate back down the bole and hide in the bark crevices. In September many specimens go back to the foliage again where they feed intermittently until the last of September. They then move down to the ground, crawl into duff where they remain until the following May. This type of migration now seems feasible for other conifer-infesting Puto, because in some California species there are known records of infestations in the duff, on the bark, and in the foliage. Areas other than the bark of conifers are infested. The foliage region is particularly well inhabited. In most instances the mealybugs are found at the bases of the needles. As far as is known, the total life cycle of these species is spent in the region of the foliage. Some species found in the foliage area are Crisicoccus pini (Kuwana), Dysmicoccus pinicolus McKenzie, D. ryani (Coquillett), Ehrhornia cupressi (Ehrhorn), Puto cupressi (Coleman), and Spilococcus implicatus Ferris (see col. pi. XXIV). Root infestations on coniferous trees are as yet unknown. It seems logical to assume, however, that with more investigation and survey mealybugs will ultimately be discovered in the root zone.

ECOLOGY Ferns Fern-inhabiting mealybugs are not at all common. Rhizoecus pritchardi McKenzie is, however, a species which is often found on the roots of maidenhair fern and was, in fact, originally described from that host. It is, however, not restricted to ferns and is perhaps more commonly associated with roots of African violet. Several other mealybugs are also found on, although not restricted to, various types of ferns in nurseries. Perhaps the most common species in this category is Pseudococcus longispinus (Targioni Tozzetti). Beardsley (1957) described several species of Pedronia collected in Hawaii which are restricted to ferns. Inanimate Objects Occasionally mealybugs may be found in association with nonbiological objects, especially during the winter period. One such area mentioned previously is under rocks (see col. pi. V). Exactly why some mealybugs are found under rocks is not clear, but it is felt that the insects are either feeding on roots beneath the rocks, are carried to the rocks by ants, or have been attracted to the cracks and crevices in the rocks as overwintering areas. Misericoccus arenarius (Doane and Steinweden) is a species which is often collected under rocks and is always associated with grass roots which come in contact with the rocks. It is interesting to note that on several occasions when this mealybug was collected on rock surfaces, only a very small number of specimens could be found on the grass roots not associated with the rocks. It was felt that perhaps this was due to collecting techniques, so Berlese samples were taken of soil under rocks and away from rocks. When these samples were compared, there were well over 75 percent more specimens found in the rock sample. Phenacoccus colemani Ehrhorn is very commonly collected in the pits and crevices under lava rocks, where it is invariably found associated with ants (col. pi. I). Many mealybug species are found under rocks, deeply imbedded in cracks. Several examples are Phenacoccus solani Ferris, P. eschscholziae McKenzie, and Heliococcus stachyos (Ehrhorn). All three of these species were found during the winter. Of the other inanimate objects which have been good areas for mealybug collecting, perhaps wooden boards have been the most important. T. R. Haig, Survey Entomologist, California State Department of Agriculture, states that at times he finds as many mealybugs under boards as in any other area. A few of the species which he has collected under boards are Chnaurococcus trifolii (Forbes), Chorizococcus rostellum (Hoke), and Phenacoccus graminosus McKenzie. MULTIPLE HOST POSITIONS It should be remembered that many mealybugs move from one position on the host to another. This may

19

depend on the time of year; it has recently been recorded that many foliage-infesting mealybugs collected in the desert in the summer are found on the crown and roots of the host in the winter. This has been shown in Phenacoccus solenopsis Tinsley and Spilococcus larreae Ferris. Host-position movement of many Discococcus species seems to best correlate to the stage of development of the mealybug. The immature stages are rather elongate and flattened and are found in the grass blade sheaths, while the rounded mature females are found on grass crowns. Movements of mealybugs from their host plant into the soil or onto the undersides of rocks have been noticed when the insect is about to shed its skin. This seems particularly true of Puto yuccae (Coquillett). Such movements also occur when mature females are about to form ovisacs and lay eggs. This has been noted in Chorizococcus polyporus McKenzie, under rocks, and Discococcus spectabilis McKenzie, in the soil. There are several species of mealybugs which are suspected of subterranean travel from host to host. Scaptococcus milleri McKenzie, herein described as new, is a species found on the unique plant Eriogonum deserticola (Polygonaceae) in the sand dunes east of Brawley, Imperial County, California. This mealybug, although often quite numerous on its host, is also found in the soil completely removed from any plant (see col. pi. XXIII). Specimens have been discovered at least ten feet away from any vegetation. Morphologically, both Scaptococcus milleri McKenzie and S. californicus McKenzie appear well adapted for digging, utilizing their exceptionally robust and heavily chitinized legs. The former species has been observed digging in the sand, the front legs alternately pulling the sand grains from the head region and passing them back to the middle pair of legs; the middle pair of legs alternately pushing the sand grains to the last pair of legs which, in turn, carry out the same process. By such a methodical system, this mealybug is capable of slow, but steady movement through the sand. Evidence that S. californicus travels in the same manner is only circumstantial, for this species has never been collected disassociated from its host. However, its front legs are proportionately quite large, and the claws normally show heavy wear. The fact that this particular species infests annual plants, which exist for not more than a month or a month and a half each year, would indicate that some movement must occur annually to new host plants; but whether this movement is subterranean is unknown. Thus, the apparent soil and sand movement of S. californicus is of paramount importance to its natural dispersal. In S. milleri, the host is a large perennial which apparently lives for many years. In this case, it would not seem as critical for the mealybug to move to new host specimens, except in an attempt to distribute itself. The dispersal of this species is, indeed, quite effective, for within its restricted sand dune locality it can be found on the roots of nearly every Eriogonum deserticola present.

20

ECOLOGY

This sort of migration suggests several questions. How do the mealybugs, as they are wandering through the soil, detect their hosts? Is this process carried out by some olfactory mechanism, or is it merely a random hit-or-miss system? No matter what the process, it is quite effective, for some mealybugs are capable of finding their annual hosts before the first shoots break the soil surface. ANT ASSOCIATIONS The classic ant-aphid mutualistic relationship has long been observed by naturalist and entomologist alike (see col. pi. V ) , but until recently few detailed studies were made on the actual benefits and factors involved in these associations. This type of relationship between ants and other insects is known to occur in a number of homopterous groups, especially in the mealybugs. In the Pseudococcidae the degree of dependence on the ants may vary from strong and almost necessary associations to weak, casual seasonal relationships. This diversity corresponds to many variable factors which all bring about a very complex association that may or may not be beneficial to both arthropods. Benefits to

Mealybugs

On collecting excursions, relationship benefits for mealybugs from association with ants are often seen. Some of the more obvious behavior of a mealybug colony is important to a basic understanding of the ant-mealybug complex. In most instances ant-tended mealybug colonies will be much larger than colonies of the same mealybug species on the same host that are not tended by ants. So outwardly the ant's presence must be of some benefit to the mealybug, either directly or indirectly. Recent study has shown that the mealybug-derived benefits are more numerous than might be expected. Way (1963) relates four possible areas in which Homoptera are benefited. Protection from Enemies This is a very important and long-realized aspect of mealybug benefit, and yet it is an area in which much controversy has arisen. Some say that the protection given homopterous insects by ants is merely accidental, since natural enemies are easily disturbed by movements of the ants. It has further been pointed out that ants are naturally hostile to any quick or obviously harmful movements around the honeydew sources (Herzig, 1938; Nixon, 1951). Other workers feel that these casual ant and mealybug habits are an oversimplification of the protective mechanisms involved. They feel that, although the above mentioned benefits are important, other factors also play major roles. Some of these factors might be the availability of other ant-food sources; whether the ant species is predaceous; or whether the parasite or predator species is well adapted to the use of ant-attended hosts. This last condition of parasite-predator adapta-

tion is often observed. Not uncommonly among an infestation of pseudococcids, another, somewhat more mobile, insect will be found. Usually it is very similar in appearance to the mealybug, and unless a trained collector is working the ladybird beetle larva will be mistaken for a mealybug. This type of larva is usually very mealy in appearance and blends in nicely with its pseudococcid host. Although ladybird beetle predators have been found feeding on mealybug colonies not attended by ants, they are often found among formicid attendants. The efficiency of this beetle predator in relation to its host depends upon the reaction which it stimulates in the ant. Even though this factor is important, it is difficult to discount the fact that the ant seems to be unable to recognize the ladybird beetle larva as a predator. There is, therefore, little doubt that the seemingly mimetic resemblence of the beetle larva to the mealybug is an aid to its more perfect predaceous habits. Another possible mealybug parasite adaptation has been noted in a species of fly larva. This larva apparently remains outside of its host's body and extracts food externally. This habit would seem to make it readily susceptible to ant attack. In order to overcome the problem, the larva is always found completely hidden beneath the mealybug, and only when the host insect is removed can the larva be seen. Other parasite-predator adaptations have been recorded, and certainly many remain to be found. Ant Constructions There are two main types of ant constructions which are important to mealybugs. The first is the actual ants' nest, and the second is the so-called carton or ant tent. The latter is not important in California, although Wheeler (1926) mentioned several instances of "carton nests" in North America. Ant-nest isolation of the mealybugs is considered quite important and has been observed on several occasions in California. In one instance, Phenacoccus artemisiae Ehrhorn was found in some of the upper chambers of a Crematogaster nest. These mealybug specimens were found at least two feet from the nearest host plant and in two instances were being transported in the mandibles of an ant. This particular collection was made in early February, which is a time of very low insect activity and high precipitation rate. The mealybugs were quiescent for the most part and were found in close contact with one another on the ceiling of ant nest chambers. The benefits derived by such an association are obvious but at least bear some mentioning. Zwolfer (1958), in his work on root aphids, suggested that the aphids are protected from moisture, fungus, and low temperatures. It seems likely also that parasite protection would be important. Fungus is apparently a major enemy of mealybugs and their relatives. On one summer collecting excursion in the extreme northeastern part of Cali-

ECOLOGY fornia, mealybugs were found after a very late series of rain storms. The amount of precipitation which had fallen previously that year was also excessively high, so that there was an overabundance of moisture in the region. A species of Eriococcus, associated with arid areas in the region, was found to be so heavily infested with fungus that it was virtually unrecognizable as a scale insect; the infestation was so excessive that not one uncontaminated specimen could be recovered. Apparently, the extra moisture made conditions perfect for fungus growth. Fungus is also a parasite of mealybugs, although infestation is not normally observed until the pseudococcid has been mounted; when examining mealybug preparations, however, fungus infestation is often seen. Ant-nest protection from a highly humid environment is therefore an important factor in mealybug welfare and is probably directly connected with protection from fungus contamination. Another mechanism which ants play in homopterous fungus protection was recorded by Pontin (1960) in connection with a species of aphid overwintering with the ant Lasius flavus. It was found that the ants constant "licking" of the overwintering eggs protected them from desiccation and "mold" contamination. It is likely that these aphids are unable to survive without the associated ant, even though summer independence may be recorded. Although the above association has never been observed in mealybugs, it is possible that a similar association does occur, for extremely close relationships between mealybugs and ants have been recorded under very similar circumstances. Protection from harsh winter conditions is perhaps the most important factor of ant-nest benefit to mealybugs in California, and there are few species, if any, which are entirely dependent upon ants for their total existence. For the most part ant tents are important in the tropical areas, where two primary types are found. The first is constructed with the silk-forming glands of the ant larvae. The adult, which has no silk gland holds a larva in its mandibles and forces the immature form to produce its silken product in the desired area. The ant genus Oecophylla is well known for this habit. The second type of tent is made of earth, "paper" formed by the ant, and leaves. Any or all of these materials may play a role in tent construction. The tents are normally built over mealybug colonies which may have as many as 1,200 individuals. Tent dimensions have been recorded up to 4.5 X 2.3 inches (Strickland, 19516). Mealybug-derived benefits are twofold. First, the tent provides some protection from direct drops of rain, although the tent itself is in no way waterproof. Second, the tent, which has only one very small entrance, is important in shielding the pseudococcids from large parasites. Apparently the mealybug parasites are unable to either find the tent entrance or push their way through. Although there are many records of tent parasitism, the rate of incidence is much lower than where no protection is afforded.

21

Direct Benefits One of the direct benefits of ant association to mealybugs is shown in relation to the production of honeydew and the subsequent contamination of the honeydew and source insect with sooty mold. Because this fungus contaminant is often the cause of mass destruction of mealybugs, certain adaptations have been made to rid the pseudococcid of the secretion. The rectum of the insect slowly fills with honeydew, and when an ant "fondles" the mealybug with its antennae, this rectal, or more often the ostiole, supply is slowly extruded for consumption by the ants. However, if ants are not present, the rectal cavity will soon fill with honeydew. When this occurs the anal valves are opened and the walls of the rectum are quickly contracted causing the honeydew secretion to fall some distance from the insect's body. In late January, 1965, an observation bearing on this matter was made on the mealybug Phenacoccus alleni McKenzie. When the pseudococcid was pulled from the soil still attached to the host roots, one specimen proceeded to squirt a small globule of honeydew from the anus to a distance of over four inches. This distance is at least 20 times the total length of the insect and shows that mealybugs are quite capable of ejecting honeydew to distances well out of range of personal contamination. Whether rapid ejection of honeydew occurs through the ostioles is unknown. When ants are in attendance, they remove the honeydew as described above, thus eliminating the problem of sooty-mold contamination. Because of this, ant-attended mealybug colonies are quite often very dense with little distance between individuals to allow for honeydew ejection. Such direct benefits as increased size of individuals attended by ants and a higher reproduction rate have been recorded as important in aphids and other homopterous insects, but they seem to be of little significance in connection with mealybugs. A final direct benefit not often realized, but perhaps a factor of importance in the understanding of the biological control of mealybugs, is that some ants actively regulate the population size of their hosts. It has been demonstrated that certain ant species will actually keep the mealybug population down to a size which they can control, thus eliminating production of honeydew. The ants regulate any excessive build-up of population by killing numbers of mealybugs; and consuming the mealybugs is considered a supplement to the protein portion of the ant diet (Way, 1963). This habit is also of importance in eliminating sooty-mold contamination in the mealybug colony. It has been observed in some heterogeneously antattended and unattended mealybug populations, with little or no parasitism occurring, that unattended colonies will get much larger than those colonies maintained by the ants. This is not always the case, but it might be of some consequence in biological control of mealybugs.

22

ECOLOGY

Ant Transport Transportation of mealybug by ants has long been known, but little has been published to substantiate this phenomenon. In California it is often possible to see ants carrying mealybugs from danger. Only in one case, however, has it been possible to watch undisturbed ants actually carrying mealybug in this fashion. In this instance an unknown species of Camponotus was observed carrying a mealybug from its host plant, directly into the ants' nest. Some instances have been noted by Strickland (1951a), who recorded actual host-to-host mealybug transport to a very small degree. Further evidence is needed to substantiate the importance of transportation in mealybug-ant associations. Benefits to Ants

The normal ant-mealybug association when observed in the field is seemingly quite simple. The ants, which may be of various genera, normally move busily from one mealybug specimen to another. When a pseudococcid is contacted, the ant begins to fondle the mealybug with its antennae just as it might fondle its own brood. On one occasion while collecting Phenacoccus echeveriae McKenzie associated with Cremotogaster sp., several revealing observations were made. The fact that the ants rested their heads on the dorsum of the mealybugs near the area of the ostioles was cause for suspicion that the ant might be receiving honeydew from the mealybug through the ostioles. This fact was soon confirmed. Mealybugs have the habit of extruding a solution from their ostioles when disturbed. This is possibly a defense mechanism. One specimen of Phenacoccus echeveria McKenzie was purposefully disturbed, and it proceeded to extrude two small globules of honeydew through its posterior ostioles. An ant then came along and within a few seconds ingested all of the extruded honeydew. Although this experiment was inconclusive, the probability of the ostioles acting as honeydew secretion mechanisms is very good. From these observations we can see that ants benefit from mealybugs in receiving honeydew from them, which is added to the ants' food supply. The amount that ants rely on honeydew for their existence varies greatly with the species involved. Some ants seem almost entirely dependent on honeydew, while others completely ignore the substance. In nutritional value, honeydew is more complete than might be expected. It may contain free amino acids, amides, proteins, many minerals, and B-vitamins (Way, 1963). The honeydew may vary greatly in its content depending on the species of mealybug, the host plant, the age of plant, the part of plant upon which mealybugs are feeding, and the length of time that the insect feeds. Normally, a complete diet of honeydew will not compensate for a protein deficiency in the ant, and supplementary protein must be added to correct the situation. A second benefit to ants from mealybugs is one of

much less importance. It is the fact that ants may use mealybug waxes for some nutritive value. There are several records of ants keeping pseudococcids wax free, but it is still unknown whether this is an adaptation to aid in procuring honeydew, or whether the normally indigestible wax is actually used in the formicids' diet. A third and final benefit has been mentioned before and deals with the use of associated mealybugs as a source of protein. Although predaceous nonhoneydew-consuming ants are not known to attend mealybugs just for protein, some ants, which have both honeydew-consuming and predaceous habits, will occasionally kill mealybugs and use them for protein. Strange Aspects of Ant-Mealybug

Associations

Exceedingly unusual observations were made by Bunzli (1935) in relation to an association between the mealybug Neorhizoecus coffeae (Laing) and the ant Acropyga paramaribensis. The eggs and immature mealybugs were kept in chambers with eggs and larvae of the ant. The immature female mealybugs apparently served as honeydew sources! When the pseudococcid matured, it no longer produced honeydew and was then transported by the ant workers to a separate mealybug chamber. The eggs that are laid by the mature females are then carried back to the brood chambers. This association is not too unusual. The extraordinary part of the relationship is the fact that the winged virgin female ants, when leaving the nest on their nuptial flight, always carry in their mandibles a fertilized female mealybug. This mealybug will soon be the beginning of the honeydew source of a newly formed Acropyga nest. Another exceptional transport association is described by Reyne (1954) in Java. In this case the mealybug Hippeococcus is especially adapted with long raptorial legs and sucker-like digitules for clinging to Dolichoderus ants. When disturbed, the highly mobile immature mealybugs climb onto the bodies of the ants and are carried into the nest where large colonies of mealybugs are maintained. In California exceptional relationships have been observed in two separate mealybug genera. In one instance, Cryptoripersia salina (Ehrhorn) was collected at Monitor Pass, Alpine Co., in association with an unidentified species of Crematogaster ant. The mealybugs, which are enclosed in a white-felted sac at maturity, were discovered when a rock was overturned. The mealybugs were all in one large chamber in the ants' nest and were in such great numbers that two handfulls could have been collected easily. The fact that no immature forms were found may confirm Bunzli's observations of a separate mealybug egg-laying chamber, but, since pursuit of the immature stages was not undertaken, confirmation cannot be certain. Once the rock had been overturned, masses of ants poured from the lower tunnels of the nest in pursuit of their honeydew "symbionts." Within a matter of 15 minutes the mealybug

ECOLOGY chamber had been emptied of its pseudococcid contents. Another unusual observation, made in Modoc County, involved an association between Phenacoccus colemani Ehrhorn and the ant Formica subpolita. This mealybug is very often collected on the underside of lava rocks in cracks and pits formed when the lava cooled. In most instances, after a mealybug infested rock was disturbed, associated ants hurriedly tore the mealybugs, waxy sacs and all, f r o m their rock habitat and carried them into cracks in the soil. T h e ants removed the mealybugs so rapidly that, in order to collect sufficient numbers of the pseudococcids, the ants had to be removed first. A final unusual record was made in Nevada but remains unconfirmed. Ants were busily tending their mealybug host in the usual fondling manner. However, in this case the mealybugs were not readily tended because apparently they were withholding their honeydew supply from the ants. T h e ants, though, had overcome this problem by butting the abdomen of the mealybug with their heads, thus causing the honeydew to flow from the mealybug ostioles. The ants then consumed the solution and repeated the process on another pseudococcid. REFERENCES Beardsley, J. W. 1957. The genus Pedronia Green in Hawaii, with descriptions of new species (Homoptera: Pseudococcidae). Proc. Hawaii. Ent. Soc. 16(2) :218-231, illus. Bunzli, G. H. 1935. Untersuchungen uber coccidophile Ameisen aus den Kaffeefeldern von Surinam. Mitt. Schweiz. Ent. Ges. 16(6-7) :453-593. Herzig, J. 1938. Ameisen und Blattlause. Ztschr. f. Angew. Ent. 24:367-435.

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Miller, D. R. 1966. Distribution patterns of selected Western North American insects.—Distribution of mealybugs on their hosts. Bull. Ent. Soc. Amer. 12(2): 111-112. Nixon, G. E. J. 1951. The association of ants with aphids and coccids. Commonwealth Inst. Ent. London 36 pp. Pontin, A. I. 1960. Observations on the keeping of aphid eggs by ants of the genus Lasius. Ent. Mon. Mag. 96:198-199. Reyne, A. 1954. Hippeococcus, a new genus of Pseudococcidae from lava with peculiar habits. Leyden. Rijks Mus. van Natuurlijke Hist. Zool. Meded. 32:233-257, illus. Strickland, A. H. 1951a. The entomology of swollen shoot of cacao.— I. The insect species involved with notes on their biology. Bull. Ent. Res. 41:725-748, illus. 1951b. The entomology of swollen shoot of cacao.— II. Bionomics and ecology of the species involved. Bull. Ent. Res. 42:65-103, illus. Washburn, R. I. 1965. Description and bionomics of a new species of Puto from Utah (Homoptera: Coccoidea: Pseudococcidae). Ann. Ent. Soc. Amer. 58(3):293-297, illus. Way, M. I. 1963. Mutualism between ants and honeydew producing Homoptera. Ann. Rev. Ent. 8:307344. Wheeler, W. M. 1926. Ants, their structure, development and behavior. Columbia University Press, New York. 663 pp., illus. Zwölf er, H. 1958. Zur systematik, biologie und okologie unterirdisch lebender Aphiden (Anoeciinae, Tetraneurini, Pemphigini und Fordinae). Ztschr. f. Angew Ent. 43:1-52.

BIOLOGY Life History, Annual Successions

LIFE HISTORY

to other hosts or at least to diverse areas of the parent host.

The life cycles of most mealybugs are not known. Some work has been conducted on the more common mealybugs, but studies on the more unusual species are almost nonexistent. A few generalizations may be made derived primarily from work done on the following species: Dysmicoccus brevipes (Cockerell), Phenacoccus gossypii Townsend and Cockerell, Planococcus citri (Risso), Puto sandini Washburn, and Spilococcus cactearum McKenzie.

Second Instar

Oviposition

Eggs are laid by the adult female among the filamentous secretion of the ovisac formed by pores on the adult's body. As many as 474 eggs have been laid by one female, but the number of eggs varies according to the mealybug species (Meyers, 1932) and the host species (Manichote, 1958, unpublished thesis). Dysmicoccus brevipes (Cockerell) bears live young, producing as many as 908 crawlers (Ito, 1938). These ovoviviparous females produce little or no ovisac and apparently shield the young for a short time by covering them with their abdomen. This character is particularly true of Puto, a genus which, for the most part, is ovoviviparous. The ovisacs are somewhat variable in their structure depending upon the species of mealybug. Several types of ovisacs have been recorded, varying from one covering the entire female body, to one covering only half or perhaps the last two abdominal segments, to one which is entirely ventral. First Instar

Nymph

The second instar numph emerges through a slit exit in the medio-dorsum of the head and thorax of the first instar skin. The legs and antennae are still proportionally larger in relation to the body than in the adult, and very often there are fewer antennal segments than in the adult. During this stage it is possible to differentiate the sexes, for by the end of this instar the males will have produced a filamentous sac or cocoon over their bodies (Ito 1938; MacGillivray 1921). It has been pointed out also by MacGillivray that the rostrum of the male is lost after the first molt, making recognition of the second instar male quite simple. Third Instar

Nymph

The second instar skin is shed in the same manner as the first, producing the third instar nymph. In female specimens this instar begins to take the normal shape of the adult. The legs and antennae are approximately the same in proportion to the body size as compared to the adult. The antennal segments are normally, but not always, the same in number as in mature forms. In the male the second skin is shoved to the posterior portion of the cocoon. The third instar male is called the prepupa and is usually much smaller and more elongate than the third instar female. It differs also in having wing pads, no rostellum, and more antennal segments.

Nymph

Adult

After approximately 3-16 days, the eggs hatch into first instar nymphs or crawlers. This stage usually has proportionally very large legs and antennae as compared with other instars, and very often there are fewer antennal segments than in the adult. This crawler is often quite mobile and usually migrates

Female

The fourth instar female emerges in the usual manner to become an adult. She can be distinguished from all of the nymphal instars by the presence of a vulva, and she is often very large and distorted from the eggs or nymphs which she carries in her 24

BIOLOGY abdomen. In appearance she has various types of pores on her body. The circular multilocular and pentagonal quinquelocular pores are believed to function in the production of the filamentous ovisac, whereas the normally more numerous trilocular pores function in the production of the white powdery secretion so characteristic of mealybugs (Ferris, 1950). The exact function of any of these pores has yet to be confirmed, and the precise morphological structure of the various types of tubular ducts is still unknown. In certain mealybug species, such as Ferrisia virgata (Cockerell), it is believed that the enlarged tubular ducts produce thin crystalline-like rods, but the importance of the smaller tubular ducts remains a mystery. The setae of the cerarii apparently are important in supporting the filaments. Also, on most adult female mealybugs, two pairs of dorsal cavities or ostioles, as they are called, can be located submarginally. The posterior pair is on the seventh abdominal segment, and the anterior pair is apparently on the head (Ferris, 1950). As has been mentioned previously, the ostioles, when irritated, seem to function in the secretion of globules of body fluid, perhaps acting in a defensive mechanism (Ferris, 1950). It is also believed that these dorsal openings function in the exit of honeydew for consumption by ants. The filaments which may be seen so often on the lateral margins of many mealybugs are apparently produced by the clusters of trilocular pores surrounding the cerarii. Other structures of the adult female are the antennae, eyes, mouthparts, legs, spiracles, vulva, circulus, anal ring, anal lobes, body setae and cerarii. The antennae of mealybugs are filiform, with the terminal segments characteristically longer and wider than the preceding ones. There is little noticeable tapering of the antennae from the first segment to the last. Antennal segmentation varies in number from two in some species of Antonina to nine in certain Phenacoccus species. Antennal shape is important in field identification of some subterranean mealybugs. The unusually short geniculate antennae of Rhizoecus, Pygmaeococcus, and Geococcus are easily recognized, particularly when compared to the normal straight antennae of most other genera, and makes their field identification quite simple. There are often some enlarged setae on the apical two antennal segments which apparently act as special sensory setae. These setae differ from the other antennal setae in being larger in diameter and more rounded at the tip. The eyes of mealybugs are compound but, according to Ferris (1950, p. 6 ) , they "are reduced to the equivalent of a single facet, which is borne upon a sclerotized base." Although these eyes are considered compound, it is believed that they function only in distinguishing between light and dark. The legs of mealybugs are often of taxonomic significance, as discussed previously, and the color of these structures may be of some importance in field identification. Although this idea is a relatively new

25

one, certain genera have been recorded as having a characteristic leg color. For example, the legs of Rhizoecus and Misericoccus are white, those of many species of Phenacoccus and Spilococcus are red, while those of Puto are very dark brown or almost black in color. Leg size is also quite variable: Antoninoides have very small, reduced legs; Discococcus have slightly larger legs; Phenacoccus, Spilococcus, Chorizococcus, and others bear legs of proportionately normal size; and numerous Puto species bear very enlarged and robust legs. The adult females of Antonina are totally without legs (apodous). On each surface of the posterior part of the trochanter there are normally two, sometimes three, small clear spots thought by Ferris (1950) to be sensory in function. These were found by DeLotto (1964) to be one of the main taxonomic characteristics of Lenania DeLotto. On the tarsal segments and on the claw, mealybugs have normally a pair of spatulate setae, but these are sometimes absent. The use of these setae is not known, but it seems logical to assume that they in some way function in clinging to the substrate. The vulva, which is always present in the adult female, is the genital opening and functions in copulation and egg laying. It is interesting to note that in some instances more than one male is able to mate with a single female at one time (Manichote, 1958; Nelson-Rees, 1959). The anal rings are often of various shapes in different mealybugs, but they all function in waste elimination. As was mentioned previously, these wastes, which are in many instances detrimental to the mealybug, are propelled long distances from the body by rapid contraction of the walls of the anal cavity. It is possible also that the anal ring serves as an exit for honeydew intended for ant consumption, and that the long setae of the ring serve as standards to hold the honeydew secretion before it is consumed by the ant. Fourth Instar

Male

The fourth instar of the male is produced in a cocoon and is known as the pupa. The third shed skin is again shoved to the back of the cocoon. This instar differs from those previously mentioned in having longer wing pads, more antennal segments (normally the same as the adult), and three pairs of eyes. At a time between this instar and the next, a break in the posterior part of the cocoon is formed and the fourth shed skin is pushed outside. Fifth Instar Male

(Adult)

The fifth instar male adult is very complex morphologically, and only the easily recognized characters are discussed here. There are two or three known primary types of dermal pores occurring on male pseudococcids (Theron, 1958). The first type is normally clustered in a circular group on the posterior lateral margin of each side of the ninth abdominal segment. In Phenacoccus gossypii Townsend and Cockerell, there is, in addition to a pair of pore

26

BIOLOGY

clusters on the ninth abdominal segment, a pair on the eighth segment. These pore groupings apparently form the wax of the long caudal filaments. The pores which make up these clusters, according to Beardsley (1960), are circular with "a five-pointed starshaped lumen," which he calls "stellate pores." It is interesting to note that most of the stellate pore clusters also have in association with them long slender setae, which apparently act as central bases for the wax of the caudal filaments. It is felt that the long anal lobe setae of the females might also function in this manner. The male caudal filaments of Puto arctostaphyli Ferris and Puto decorosus McKenzie, herein described as new, show a seta in the middle of the long waxy filaments. In Phenacoccus gossypii Townsend and Cockerell and Pseudococcus longispinus (Targioni Tozzetti), the above character was observed in the females. Other types of pores have been found on the males and are scattered over the body surface. Beardsley (1960) merely calls these "dermal disc pores" and names the number of loculi which they contain. This may vary from three to five in number with four being normal. It is further pointed out that these structures are by no means flat, but rather are recessed into the derm with several small protrusions from the central part of the loculi. The exact function of these pores is completely unknown, but it is reasonable to suspect that, along with a secretory mechanism, they may also have a sensory function. Body setae are also characteristic in the male. Although the normal lanceolate setae of the female are present, so also are the "thick, finger-like digitiform setae" described by Beardsley. The latter setae are normally the most common of the two types; they predominate on both body surfaces, the legs, and especially the antennae. The "digitiform" setae, which are presumably sensory in function, are very similar to those on the last two segments of the females' antenna. The position of the ostioles and the exact designation of abdominal segments is presently a matter of some disagreement. For simplicity sake the view of Ferris (1950) shall be maintained, although it should be understood that I do not feel that one theory is better than the other. In accordance with Ferris' interpretation of the male segmentation, one pair of dorsal ostioles is present on the submarginal portion of the seventh abdominal segment and corresponds to the posterior pair of ostioles on the female. The antennae of the adult male are normally tensegmented, but there are occasionally species which have nine-segmented antennae, such as Palmicola palmarum (Ehrhorn). The male has six eyes—a ventral pair, a lateral pair, and a dorsal pair. According to Beardsley (1960), the eyes of most males are dark red, with the smaller lateral pair protruding more than the other two. The mouthparts of the male are almost nonexistent and are completely nonfunctional. All that remains of the mouth is a small circular opening found

at the posterior margin of the ventral part of the head. There is normally one pair of wings which are developed from the mesothorax. These wings possess two longitudinal veins, the anterior one the radius, and the posterior one the media. There is also a complex of minute basal veins which Giliomee (1961) calls the "costal complex of wing veins." There are three forms of males, the first form has fully developed wings (macropterous); the second form has wings present but reduced and nonfunctional (brachypterous); and the third form is wingless (apterous). Examples of the first form are Phenacoccus gossypii Townsend and Cockerell, Planococcus citri (Risso), Puto laticribellum McKenzie, and Saccharicoccus sacchari (Cockerell); of the second form Palmicola palmarum (Ehrhorn); and of the third form, Puto ambiguus (Fullaway), P. echinatus McKenzie, and P. pacificus McKenzie, herein described as new. On the metathorax a pair of halteres is present. They are slender projections adorned with one or more setae at their tips. The normal number of apical setae is one, but Beardsley (1962a) found four on each haltere in Puto yuccae (Coquillett). He further stated that the apical setae, whether four or more in number, are recurved in such a manner as to hook into a circular pocket in the posterior part of the forewing. There are two pairs of spiracles in the male just as in the female, but in the case of the male there is often a much more strongly developed peritreme. The legs of most males are quite similar to those of the females, although they are usually proportionately much longer and more slender. The tarsus of the male has two segments rather than the normal single segment of the female, although in most cases the additional segment is quite small and appears as a slightly sclerotized ring. The digitules of the tarsal claws of most males differ from the spatulate type of the female in being slender and setiform. The claw is normally without a denticle. The external male genitalia is relatively simple, consisting of a large penial sheath, a conspicuous vertical slit, and the penis or aedeagus. The penial sheath is normally large and sclerotized with a posterior apical projection. This projection may be of various shapes but is usually broadly oval. On the ventral surface of the sheath there is a noticeable slit from which often protrudes the aedeagus. This structure is normally tube-shaped and curves downward. The penial sheath is apically of various shapes, from broad as in Pseudococcus fragilis Brain to very sharp and pointed as in P. longispinus (Targioni Tozzetti) (Giliomee, 1961). Beardsley (1962b) gave a description of the male of Puto yuccae (Coquillett). This species, as well as many others of the Puto group, is quite different from the "normal" pseudococcid male. The primary differences noted were eight pairs of eyes, four setae on each halter, a toothed and bifurcate aedeagus, and a denticle on the claw.

BIOLOGY Parthenogenesis Male mealybugs are quite common and are very often essential for reproduction. The reason that they are overlooked is that they live for only a few days and are normally, although not always, quite small. Parthenogenesis does occur in coccids, but this subject is quite complex. Various types of parthenogenesis occur along with functional hermaphroditism. There seems also to be races of the same species which are either entirely parthenogenetic or are entirely bisexual (Hughes-Schrader, 1925). Further details on sex ratio and unisexual (parthenogenetic) races will be considered in the chapter on cytology. Instars and Generations The numbers of instars which mealybugs go through is four for the female and five for the male. Although Uichano and Villanueva (1932) described Saccharicoccus sacchari (Cockerell) as having seven instars in both sexes, Beardsley (1962a) found in Hawaii, at least, that this mealybug has only the normal four female instars and five male instars. The number of generations is quite variable in the Pseudococcidae. Myers (1932), in his studies on Planococcus citri (Risso) and Phenacoccus gossypii Townsend and Cockerell, found it possible to carry both of these species through eight life cycles in approximately one year. In most of the Puto species, however, only one generation occurs annually, and in Puto sandini Washburn, a high altitude species, there is one generation every four years (Washburn, 1965). Again, we see a wide variation throughout this extremely large family Pseudococcidae. Succession in Mealybugs As previously mentioned, my assistant and I have devoted the past few years to rather extensive mealybug collecting. During the collecting seasons a trend of succession, or species replacement, was noted. In early spring of the year certain species are more prominent than others. As spring passes to summer, other species replace the once common spring forms, and the summer species are in turn replaced by the fall and winter species. This trend may occur on a single host species, or it may be apparent on all hosts in a given habitat. This type of species replacement seems quite logical. It has been previously noted that mealybugs are extremely sensitive to varying amounts of moisture and humidity, and, as the seasons change from spring to fall, the environment becomes less favorable for some mealybugs and more favorable for others. For certain species the prominence of specific host plants at rather definite times of the year is an important factor, and, if micro-environmental conditions around the hosts are not favorable to the mealybug, it will not survive.

27

REFERENCES Beardsley, J. W. 1960. A preliminary study of the males of some Hawaiian mealybugs (Homoptera: Pseudococcidae). Proc. Hawaii Ent. Soc. 17(2):199-243, illus. 1962a. Notes on the biology of the pink sugar cane mealybug, Saccharicoccus sacchari (Cockerell) , in Hawaii (Homoptera: Pseudococcidae). Proc. Hawaii. Ent. Soc. 18(1) :55-59. 1962b. Descriptions and notes on male mealybugs (Homoptera: Pseudococcidae). Proc. Hawaii. Ent. Soc. 18(1) :81-98, illus. De Lotto, G. 1964. Observations on African mealybugs (Hemiptera: Coccoidea). Bull. Brit. Mus. (Nat. Hist.) Ent. 14(8) :343-397, illus. Ferris, G. F. 1950. Atlas of the scale insects of North America (series V). The Pseudococcidae (Part I). Stanford Univ. Press, Stanford. 278 pp., illus. Giliomee, J. H. 1961. Morphological and taxonomic studies on the males of three species of the genus Pseudococcus (Hemiptera: Coccoidea). Ann. Stellenbosch Univ. 36 (ser. A) 6:243-296, illus. Hughes-Schrader, S. 1925. Cytology of hermaphroditism in leery a purchasi (Coccidae). Ztschr. f. Zellforsch. und Mikros. Anat. 2:264-292, illus. Ito, K. 1938. Studies on the life history of the pineapple mealybug, Pseudococcus brevipes (Ckll.). Jour. Econ. Ent. 31(2) :291-298, illus. MacGillivray, A. D. 1921. The Coccidae. Scarab, Urbana, 111. 502 pp. Manichote, P. 1958. The biology of the cactus mealybug, Spilococcus cactearum McKenzie (Homoptera: Coccoidea: Pseudococcidae) (Unpublished thesis for Master's Degree, 66 pp., illus.). Myers, L. E. 1932. Two economic greenhouse mealybugs of Mississippi. The citrus mealybug and the Mexican mealybug. Jour. Econ. Ent. 25(4):891896, illus. Nelson-Rees, W. A. 1959. Triple coitus in the mealybug, Planococcus citri (Risso). Nature 183:479. Theron, J. G. 1958. Comparative studies on the morphology of male scale insects (Hemiptera: Coccoidea) Ann. Stellenbosch Univ. 34 (sec. A) 1:1-71, illus. Uichango, L. B. and F. E. Villanueva 1932. Biology of the pink mealybug of sugar cane, Trionymus sacchari (Cockerell), in the Philippines. Philippine Agr. 21:205-276, illus.

CYTOLOGY

early development of the male embryos. This set does not undergo the usual changes during a division cycle, but maintains a compact and densely staining form. This set can thus be recognized in the non-dividing nucleus; sometimes its chromosomes are individually distinguishable, but frequently all the members of a paternal set come together to form a dense, lens-shaped mass at one side of the nucleus (Figure 1). The presence of the dense, dark staining bodies in many or all of the nuclei makes it possible to tell at a glance whether or not males occur and whether the species is, therefore, bisexual. In the primitive genus of mealybugs, Puto, the females have two X chromosomes, and the males only one. This is an orthodox system, found in many insects. It resembles the better known X X ( 2 ) - X Y ( $ ) system and was undoubtedly derived from it by evolutionary degeneration and final loss of the Y chromosome. We can therefore symbolize the system as X X ( 2 ) X O ( 3 ) . Sex is determined genetically. All eggs have an X chromosome. Half the sperm carry an X chromosome and yield X X daughters. The other half carry no sex chromosome (the " O " sperm) and yield X O sons (Table 1). In the lecanoid chromosome system which characterizes the majority of the mealybugs, there are no sex chromosomes and only one chromosome type of egg and one chromosome type of sperm are found (see reviews of Hughes-Schrader, 1948; and Brown and Nur, 1964). By some unknown mechanism, a decision is made inside the egg as to whether the embryo is to be male or female. If it is to be male, the paternal set of chromosomes brought in by the sperm undergo the condensation process (heterochromatization) already described. The condensed chromosomes undergo normal divisions and are maintained as such in most, but not all, of the nuclei of the developing male. The reduction divisions of the male in the lecanoid

For many years the mealybugs have been known to have unusual systems of inheritance. One early approach, that of Schrader (1921; see review of Hughes-Schrader, 1948), was a direct examination of the unorthodox behavior of the chromosomes. Other studies, such as that of James (1937), showed that the sex ratio is extremely variable, much more so than would be expected if a typical sex-determining mechanism were in operation. James noted that the sex ratio varied from 1 i : l 2 to 1 S : 5 2 among the five species he studied. Furthermore, variation in sex ratio of the progeny of individual females was quite marked in one species, where it ranged from 1 3 : 6 2 to 4 3 : 1 2 . James also showed that aging the mother prior to mating markedly altered the sex ratio in favor of the males, and more recently Nelson-Rees (1960) discovered that the sex-ratio varies during oviposition and will be different for the embryos produced during the first few days from those produced later. The complex puzzle of the mechanism of sex determination was finally resolved by Nur in work to be discussed later. It should be noted here, however, that as a result of the variable sex ratio, the tendency for the males to hide, and the very short life span of the adult male, the field worker can be confused about the significance of males in a given species or single colony. Because many species of coccids are unisexual, consisting only of females producing daughters parthenogenetically (thelytoky), confusions of unisexual and bisexual species have often occurred. Furthermore, some species of coccids have both sexual and parthenogenetic races (HughesSchrader, 1948). GENETIC SYSTEMS The most striking aspect of the mealybug, the "lecanoid" chromosome system, is the change in the paternal chromosome set which occurs during the 28

29

CYTOLOGY

Fig. 1. Photomicrographs of nuclei not in division. A. After the earliest stages of embryonic development, the heterochromatic set appears prominently in nuclei of the males. The individual chromosomes may be obvious or may all be gathered together in an amorphous mass. B. The nuclei of very young embryos, and of females, do not show such development of heterochromatin. system are also unusual (Figure 2). There are two divisions. During the first of these, both the paternal and maternal sets simply divide as they do in mitotic cell divisions during development. During the second division, the paternal and maternal sets are segregated on opposite sides of the cell. Only the maternal set forms a sperm. The paternal set degenerates. The male mealybug thus transmits to his offspring only the genes and chromosomes which he received from his mother. The net effect is exactly the same as in the well-known example of the honey bee. Here the male develops from an unfertilized egg, contains only the maternal chromosomes, and transmits them to his offspring. It is an intriguing aspect of the evolution of chromosome systems that mealybugs have a complex mechanism to do what honey bees do so simply.

curs also in certain coccids but not in the mealybug family or its evolutionary descendants. Parthenogenesis is frequent in the latter groups but is restricted to the production of daughters (thelytoky). There are two main types of parthenogenesis. In one, an ordinary division of the chromosomes is substituted for the reduction divisions. The egg thus has a full set of chromosomes and can proceed directly with development. In the other type, reduction divisions occur so that there is only one set of chromosomes in the egg nucleus. The egg can combine with one of the other products of the reduction division; it will then have the full quota of chromosomes necessary to proceed with development. Or the egg may divide to give two daughter nuclei; these then come back together to produce one nucleus with the necessary number (Figure 3).

Parthenogenesis

Sex

In some insects, such as the honey bee and other Hymenoptera, the males are produced parthenogenetically from unfertilized eggs. This mechanism oc-

Determination

Oddly enough, it was a study of a parthenogenetic species by Nur (1963) which gave us conclusive evidence of the nature of sex determination in the

TABLE 1 CYTOLOGY OF SOME MEALYBUGS

Species Puto sp. Pseudococcus longispinus Pseudococcus obscurus Pseudococcus straussiae Antonina graminis Clavicoccus tribulus Dysmicoccus brevipes Phenacoccus acericola Planococcus citri Rhizoecus falcifer Saccharicoccus sacchari Trionymus rostellum a b

Chromosome Number 9 ê 14 8 10 38 16 14 10 10 12 10 12 10 10

13 8 10 (38)b 14 10 12 10 12 10 10

System XX-XO lecanoid lecanoid lecanoid parthenogenetic lecanoid lecanoid parthenogenetic lecanoid lecanoid lecanoid lecanoid lecanoid

1—Hughes-Schrader, 1948; 2—Nur, 1962; 3—Brown, Beardsley, DeLotto, and Nur (unpublished). Chromosomes not counted, but typical male heterochromatization present in some embryos.

Reference a 1 3 2 3 3 3 3 3 1 1 3 3 3

30

CYTOLOGY

//l\ 8 0 « J

> A B C Fig. 2. Diagram of spermatogenesis in the mealybug (lecanoid) chromosome system. The maternal, euchromatic set (light) and the paternal, heterochromatic set (dark) are here indicated by only two chromosomes each. During the first division, both types divide equationally (A and B). During the second division, the two types are segregated (C). Only the euchromatic derivatives form sperm; the heterochromatic set forms a dense-staining residue which soon degenerates (D). mealybugs. Although Nur studied the species in another family, they have the mealybug, or lecanoid, chromosome system. The parthenogenetic species studied produces mostly females by the second method of parthenogenesis described above, but in a small percentage of cases, one of the two daughter nuclei is apparently mistaken for a sperm, because the typical condensation process appears later. This exceptional behavior is quite rare and should not be a serious source of error in analysis of a species for bisexuality. Although no sperm were present in this case, parthenogenetic females of other species have been known to mate and be inseminated. Chromosome

Number

Most of the common mealybugs have 10 chromosomes, 5 from each parent. In the males, the 5 of the paternal set show the condensation process, while the 5 of the maternal set do not. This is the basic number for the family and occurs so commonly that chromosome number studies are useful in distinguishing species only in a few instances. In many divisions, the formation of a nuclear organelle, the nucleolus, has produced a long gap in one of the chromosomes; if this situation is not recognized, an extra chromosome may be erroneously counted. Mealybug chromosomes, like those of other Hemiptera, do not have localized centromeres dividing the chromosomes into two arms; however, flexures occur which sometimes resemble those commonly seen at the centromere in other organisms. The mealybugs often make large cells from the nuclei (polar bodies) left over from the reduction divisions which produced the egg. If the number of chromosomes per set is five, these large cells will start out with 15 chromosomes, change to 25, and then to higher chromosome numbers (Figure 3). In parthenogenetic forms, the original number may be 10, but it will soon escalate also. The large cells produced in this fashion continue to live in the embryo and usually, but not always, house the symbionts. The cells housing the symbionts, whether the large cells just referred to or others differently derived, are called mycetocytes. The symbionts are

yeast-like in appearance and are apparently necessary for the insect's metabolism. Symbionts occur in both males and females; in the nuclei of the symbiont cells in the males there are usually no condensed chromosomes. TECHNIQUE Cytological methods provide a quick and convenient way for the field worker to check on the sexuality of the species and often to determine the chromosome number. The easiest technique is to use gravid females, and the best time to collect these is just when they are commencing to "fuzz out" to build an egg sac,

PO PO

,7 ^ f' PB1

i7f"5>» / n , \ I \ \ f r i . ; EGG

PBl

I .S; —" \>N

' r

• •

K v»-

r" 1 . .'

••

* * .

•

—

—

Fig. 66. Heliococcus atriplicis McKenzie, collected on saltbush, A triplex (Chenopodiaceae), at Brawley, Imperial County, California.

lentijormis

185

186

TAXONOMY

OF

SPECIES

Recognition Characters. Adult females, mounted, 2.80 to 3.60 mm long, 1.60 to 2.30 mm wide; body shape broadly oval. Dorsum with only 2 pairs of cerarii. Anal lobe cerarius somewhat sclerotized, with 2 small, slightly lanceolate, conical setae, a few small and inconspicuous auxiliary setae, and 4 or 5 enlarged tubular ducts, but without noticeable clustering of trilocular pores. Penultimate cerarius somewhat sclerotized, with 2 small, slightly lanceolate, conical setae, but without auxiliary setae, tubular ducts, or clustering of trilocular pores. Trilocular pores generally scattered over dorsum. Characteristic enlarged tubular ducts of 2 markedly different sizes: the larger with duct prominence short and broad, having 3 or 4 small setae arising from its base; the smaller ducts relatively more slender, occasionally with only 1 seta. Larger ducts numerous on dorsum, arranged in submarginal band from apex of abdomen to head and in an irregular series across certain abdominal segments, usually sixth and eighth; the smaller ones still more numerous, occurring in a definite submarginal band from eighth abdominal segment to tip of head and in irregular rows across abdominal segments, thorax, and head. Body setae all very small, distributed over entire body surface. Anal ring apical, with the distinctive structure of this genus; each of its 6 setae longer than diameter of ring. Venter with a small, irregular sclerotization at base of anal lobe seta. Not more than 5 multilocular disk pores, immediately around the vulva. Quinquelocular pores quite numerous from apex of abdomen to head. Trilocular pores relatively sparse, noticeable along body margin but lacking in some areas in midabdominal and sternal regions. Tubular ducts, similar to the smaller ones on dorsum, in some numbers along submarginal areas. Body setae slender, generally much longer than those on dorsum. Circulus proportionately large, transversely oval, anterior margin often slightly produced laterally. Legs well formed, with a noticeable denticle on claw. Antennae 9-segmented. Notes. This species is different from all other North American components of Heliococcus in possessing only 2 pairs of cerarii—an anal lobe and penultimate pair—whereas the other known species have the normal 18 pairs of cerarii. California Records (Map 62) Imperial Co.: Brawley, XI-28-62, on Atriplex lentiformis (V. D. Roth, CDA).

Heliococcus stachyos (Ehrhorn) Betony Mealybug (Figure 67) stachyos Ehrhorn, 1900. Canad. Ent.

Phenacoccus 32:313. Phenacoccus stachyos Ehrhorn, Hollinger, 1918. Ibid. 50:23-24. Phenacoccus stachyos Ehrhorn, Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., pp. 59-60, illus.

Heliococcus stachyos (Ehrhorn), Ferris, 1950. Atlas of scale insects of North America, Ser. V: 101— 102, illus. Heliococcus stachyos (Ehrhorn), McKenzie, 1960. Hilgardia 29 ( 1 5 ) : 707. Type Locality and Host. San Francisquito Canyon, near May field (Stanford University), Santa Clara County, California, on Betony, Stachys bullata (Labiatae). North American Distribution. Known only from California. Additional Hosts. Some additional hosts of this mealybug are as follows: Adenostoma fasciculatum (Rosaceae), Arctostaphylos sp. (Ericaceae), Diplacus aurantiacus (Scrophulariaceae), Monardella sp. (Labiatae), Rhus diversiloba (Anacardiaceae), and Solanum umbellijerum (Solanaceae). Unpublished host records include: Acer platanoides (Aceraceae), Catalpa sp. (Bignoniaceae), Encelia farinosa (Compositae), Eriogonum inflatum (roots), Eriogonum umbellatum var. polyanthum (roots) (Polygonaceae), Populus sp. (Salicaceae), Prunus sp. (Rosaceae), Thermopsis (?) sp. (Leguminosae), and Ulmus sp. (Ulmaceae). External Features and Habitat. In life this mealybug is reported as covered with a mealy secretion from which there arise many delicate, glassy threads of wax. The lateral and caudal tassels are very short. The species is oviparous and does not form a definite ovisac. The mealybug may be found on the twigs, branches or roots of its host. Recognition Characters. Adult female, mounted, 1.95 to 3.50 mm long, 1.10 to 1.65 mm wide; body shape broadly oval. Dorsum with 18 pairs of cerarii. Anal lobe cerarius

TAXONOMY

OF

SPECIES

Fig. 67. Heliococcus stachyos (Ehrhorn), collected on Betony, Stachys bullata (Labiatae), at San Francisquito Canyon, near Mayfield (Stanford University), Santa Clara County, California. (Illustration prepared from specimens collected on Rhus diversiloba [Anacardiaceae], at Stevens Creek, near Stanford University, Santa Clara County, California.)

187

188

TAXONOMY

OF

SPECIES

with 2 rather small, slightly lanceolate setae, 3 enlarged tubular ducts, several short auxiliary setae, few associated trilocular pores. Remaining cerarii with 2 setae, smaller than those of anal lobes, 4-5 trilocular pores, not at all elevated upon prominences. Dorsum with trilocular pores distributed very sparsely. Enlarged tubular ducts on dorsum of 2 sizes, infrequently smaller sized ones absent, larger ones much predominating, on most segments a single larger duct on each side near median line, 1-3 such ducts in marginal area near each cerarius. A few smaller size ducts occur especially along margin. Dorsal body setae few and very small. Anal ring apical, distinctive for genus, bearing 6 setae, each longer than diameter of ring itself. Venter with multilocular disk pores extremely few or wanting—none observed in which total of these structures is more than 8—these situated immediately about vulva. Trilocular pores very sparsely distributed on venter, noticeable clear areas showing especially in midregion of abdomen and thorax. Quinquelocular pores present in midregion from vulva to head, these sparse. Small size tubular ducts, similar to those on dorsum, situated along body margins, absent elsewhere. Body setae few, for the most part longer than those on dorsum. Circulus present, small, oval. Anal lobe seta with small, elongate, sclerotized area. Legs well developed; claw with distinct denticle or tooth. Antennae 9-segmented, slender. Notes. This species seems rather closely related to Heliococcus atriplicis McKenzie, but differs from the latter in possessing 18 pairs of cerarii as compared to only 2 pairs of cerarii in the species last mentioned. Heliococcus stachyos is also similar to an eastern species, H. osborni Ehrhorn, but the more numerous dorsal enlarged tubular ducts, and fewer smaller ducts, readily separates it from that form which has only a few dorsal enlarged ducts and more numerous ducts of the smaller size. California Records (Map 63) Alpine Co.: Monitor Pass, VIII-2-64, under rocks (D. R. Miller and J. A. Froebe, UCD). El Dorado Co.: Green Valley, V-30-31, on Ceanothus sp. (H.H. Keifer, CDA); 5 miles north Smith Flat, South Fork American River, VII-12-62, on Arctostaphylos sp. (L. A. Stange, UCD). Fresno Co.: 4 miles northeast Auberry, Jose Basin Road, VIII-3-56, in "soil sample" (R. O. Schuster, UCD); Balch Camp, VIII-13-56, in "soil sample" (R. O. Schuster, UCD); Coalinga, 111-21-58, on

Eriogonum inflatum (roots) (H. L. Wilson, CDA). Mariposa Co. : 2 miles west Coulterville, V-28-65, on Adenostoma fasciculatum (R. P. Allen, CDA); 1.5 miles west Mt. Bullion (formerly known as Princetown), IV-29-63, on Adenostoma fasciculatum (R. P. Allen, CDA). San Bernardino Co.: 10 miles southwest Hesperia, IV-17-65, on Boraginaceae (foliage) (D. R. and J. F. Miller, UCD). San Mateo Co.: Woodside, III—28-17, on Stachys bullata (G. F. Ferris, UCD). Santa Clara Co.: Parmanente Creek, V-16-17, on Solanum umbelliferum (G. F. Ferris, UCD); Stevens Creek, VIII-26-16, on Rhus diversiloba (G. F. Ferris, UCD). Santa Cruz Co.: Big Basin, VI-10-17, on Stachys sp. (G. F. Ferris, UCD). Sierra Co.: 3 miles northeast Sierraville, VII-664, on Mentzelia sp. (crown) (D. R. Miller, UCD). Siskiyou Co. : Valentine Cave, Lava Beds National Monument, VI-29-63, on Eriogonum umbellatum var. polyanthum (roots) (D. R. Miller, UCD).

HETEROCOCCUS Heterococcus Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., p. 65. Genotype. Heterococcus arenae Ferris. Recognition Characters. Pseudococcidae with body form elongate elliptical to almost parallel-sided, length

FERRIS

usually more than twice width. Dorsal ostidles present. Recognizable cerarian development at most restricted to posterior 2 to 4 pairs, and cerarii, even here, not sharply defined, the association of two elongate setae, quinquelocular pores, and auxiliary

TAXONOMY

setae loose, sometimes even indistinct; additional cerarii on anterior abdominal segments, on head above eyes, more or less suggested but not clearly evident. Anal lobes at most rounded bulges. Trilocular pores absent, replaced by quinquelocular pores on both body surfaces. Multilocular disk pores present or absent. Tubular ducts of modified oral-collar type, without well-defined oral collar, on both dorsum and venter. Circulus absent. Venter with multilocular disk evident. Legs normal, a little thickened, no pores on hind coxae, tiny ones scattered on hind femora and tibiae; tarsal digitules slender, acute, not exceeding claw apex, claw digitules slender, slightly knobbed apically, definitely exceeding claw apex, claw denticle developed. Antennae characteristically 9-segmented, infrequently with 8 segments. Notes. The significant morphological features of this genus are the presence of quinquelocular pores on dorsum and venter which replace the normal trilocular pores; the 9-segmented antennae; and the claw usually with denticle or tooth present on planter surface. To the genus Heterococcus there are now assigned 3 species for North America, 2 of which occur in California. The genus appears to be Holarctic.

KEY Species of Heterococcus in North America: Adult Females

1.

2(1).

Anal lobe cerarii well defined and developed, each with 2 stout, conical setae 2 Anal lobe cerarii not clearly defined, each represented by slender setae which can be considered cerarian setae only on the basis of their paired character arenae Ferris Anal ring setae normal, slender, longtapering, longer than diameter of ring itself; anal lobe setae similarly long pulverarius (Newstead) Anal ring setae stout, not equaling in length diameter of ring itself, often bifurcate apically; anal lobe setae scarcely longer than those of anal ring occidentalis Morrison

Heterococcus arenae Ferris Sand Mealybug (Figure 68) Heterococcus arenae Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., p. 65, illus. Heterococcus arenae Ferris, 1919. Ibid., p. 22. Heterococcus arenae Ferris, Morrison, 1945. Jour. Wash. Acad. Sei. 35 (2):46-48, illus. Heterococcus arenae Ferris, 1953. Atlas of scale

OF

SPECIES

189

insects of North America, Vol. VI: 364—366, illus. Type Locality and Host. Pacific Grove (sand dunes), Monterey County, California, on a bluegrass, Poa douglasii (Gramineae). North American Distribution. California and New Mexico. Additional Hosts. This mealybug is found only on members of the grass family, Gramineae. Additional grass hosts include: Agropyron sp., Aristida sp., Elymus sp., "grass", Stipa pulchra, and unknown perennial grass. External Features and Habitat. This species is apparently surrounded by a small amount of white secretion and is thinly coated with a white, waxy powder. It occurs beneath the leaf sheaths of its host. Recognition characters. Adult female, mounted, 1.40 to 3.50 mm long, 0.40 to 1.75 mm wide; form normally elongate, slender. Dorsum with anal lobes, each bearing a small cluster of slender setae, 2 of which are so arranged that they probably represent paired cerarian setae, but which are so similar to other setae (possibly auxiliary) about them that no definite distinction can be recognized. Cerarii absent anterior to anal lobes. Trilocular pores lacking, replaced by numerous quinquelocular pores beset over entire dorsum. Mingled with quinquelocular pores, relatively small number of tubular ducts without an oral collar. Body setae small, sparsely distributed. Anal ring situated at posterior apex of body; its 6 setae but little longer than longitudinal diameter of ring itself. Venter with multilocular disk pores present in small numbers on median regions of abdominal segments 7 to 10. Quinquelocular pores numerous, rather evenly beset over ventral surface of body. Tubular ducts similar to those on dorsum, distributed similarly. Body setae sparsely situated, some slightly longer than those on dorsum. Circulus absent. Legs of normal size and form for genus; posterior pair with few translucent pores on coxa, femur and distal part of tibia; claw slender, denticle present, at times obscure on plantar surface. Antennae 9-segmented. Notes. The presence of a 9-segmented antenna, tarsal claw with denticle, and replacement of the usual trilocular pores with quinquelocular pores, readily distinguishes members of Heterococcus from all other California pseudococcids. Heterococcus arenae Ferris shows considerable size variation among specimens examined, although no morphological bases is evident to separate them. This mealybug is related to Heterococcus pulverarius (Newstead) ( = H. graminicola Morrison) but differs in the absence of dorsal multilocular disk pores on abdomen, whereas in the species last mentioned these structures are definitely present in this area and on the body surface. California Records (Map 64) Alpine Co.: 3 miles north Markleeville, VIII-9-64, on Gramineae (leaf sheaths) (D. R. Miller, UCD).

190

TAXONOMY

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TAXONOMY

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SPECIES

191

Inyo Co.: Antelope Springs, VI-24-64, on Elymus triticoides (leaf sheaths) (D. R. Miller, UCD); Between Big Pine and Deep Springs (alt. about 7,000 feet), VII-1918, on "unknown perennial grass" (G. F.Ferris, UCD). Monterey Co.: Dunes near Pacific Grove, XII1-17, on Poa douglasii (G. F. Ferris, UCD). Napa Co.: 4 miles west Spanish Flat, III—1—61, on "grass" (crown) (H. L. McKenzie, UCD). Orange Co.: Modjeska Canyon, 1-31-64, on Stipa pulchra (sheaths) (D. R. Miller and J. A. Froebe, UCD); VI-12-64, on Stipa sp. (D. R. Miller and J. A. Froebe, UCD). Placer Co.: 3 miles west Applegate, VIII-8-64, on Gramineae (leaf sheaths) (D. R. Miller, UCD). Riverside Co.: 3 miles south Temecula, II-2-64, on Aristida sp. (leaf sheaths) (D. R. Miller and J. A. Froebe, UCD). San Bernardino Co.: 1.5 miles north Cajon Pass, VI-12-60, in "soil-grass association" (H. L. McKenzie, UCD). San Diego Co.: 5 miles southeast Fallbrook, V-9-64, on Gramineae (leaf sheath) (D. R. Miller, UCD).

Heterococcus graminicola Morrison, 1945. Jour. Wash. Acad. Sci. 35(2):48-53, illus. Heterococcus variabilis Schmutterer, 1958. Prague Narod. Mus. Ent. Oddeleni Sborn. Faunistickych Praci 3:17-22, illus. Heterococcus pulverarius (Newstead), Williams, 1961. Bull. Ent. Res. 51 (4):673-675, illus. Heterococcus pulverarius (Newstead), Williams, 1963. Entomologist 96:101. Type Locality and Host. Wooster, Wayne County, Ohio, on Timothy, Phleum pratense (Gramineae). North American Distribution. California, New Jersey, Oregon, Pennsylvania, Washington, and Washington, D. C. Additional Hosts. This mealybug has been found on the following Gramineae: Agropyron repens, Alopecurus sp., Arrhenatherum elatius, Bromus carinatus, Bromus sterilis (?), Elymus glaucus, Festuca rubra, Festuca spp., "grass", Holcus lanatus, Hordeum murinum, "lawn grass", Lolium perenne, Poa sp., Setaria viridis, and "undetermined grass." External Features and Habitat. According to Morrison (1945), this mealybug is pale yellow, produces some white, waxy secretion, and is lightly coated with white secretion. It is found between the leaf sheaths and stems of the host (see col. pis. I and XI). Recognition Characters. Adult female, mounted, 1.41 to 3.25 mm long, 0.53 to 1.60 mm wide; body form elongate, at least twice as wide as long. Dorsum with posterior 3 pairs of cerarii fairly plainly developed, 4th and 5th from caudal apex often suggested by a pair of recognizable cerarian setae, the 2 setae in each pair distinctly separated, but appreciable variation occurring in extent of development of all these cerarii; ocular cerarii present, represented by 3 small conical setae, accompanied by 2-3 quinquelocular pores. Anal lobe cerarius with 2 small conical setae, 15-18 loosely grouped quinquelocular pores, 3-6 stiff, short, auxiliary setae. Trilocular pores absent, replaced with numerous small, quinquelocular pores over entire dorsum. Multilocular disk pores present from eighth abdominal segment to head, concentrated mostly along body margins, distinct patches particularly noticeable in lateral areas of eighth abdominal segment and head. A very few exceedingly small tubular ducts situated along lateral margins of abdominal segments, absent elsewhere on dorsum. Dorsum beset sparingly with extremely small conical or slightly lanceolate body setae. Circulus absent. Legs well developed, posterior pair with pattern of scattered, tiny, translucent pores on femur and tibia, none on coxa; claw denticle usually quite distinct. Antennae 9-segmented.

Heterococcus pulverarius (Newstead) Bluegrass Mealybug

Notes. This species is similar to Heterococcus arenae Ferris, but differs in possessing dorsal multilocular disk pores, whereas in the species last mentioned these structures are restricted to the ventral surface of body. Williams (1963) stated the following relative to pertinent synonymy of certain species with Hetero-

(Figure 69; Color Plates I and XI) Ripersia pulveraria Newstead, 1892. Ent. Monthly Mag. 28:145. Phenacoccus nudus Green, 1926. Ibid. 62:172-173, illus.

192

TAXONOMY

OF

SPECIES

Fig. 69. Heterococcus pulverarius (Newstead), collected on "bluegrass," Poa sp. (Gramineae), east of Tulelake, Modoc County, California. (Illustration prepared from type material.)

COLOR PLATE XI. The bluegrass mealybug, Heterococcus pulverarius (Newstead), infesting perennial ryegrass, Lolium perenne (Gramineae), Davis, Yolo County, California.

TAXONOMY

coccus pulverarius (Newstead): "No specific differences have been detected between H. graminicola and H. pulverarius. The specimen of H. variabilis possesses 6-segmented antennae and very small legs and in this respect is identical with the type specimens of H. pulverarius with 6- or 7-segmented antennae. This reduction in leg size in relation to the small antennae has been mentioned by Schmutterer. "Numerous specimens have recently been studied which were kindly collected by Dr. V. F. Eastop of the British Museum (Natural History) at Keiv. All of these have either 8- or 9-segmented antennae and the leg sizes agree with those on all the previously studied British specimens with 9-segmented antennae together with specimens of H. graminicola. Although the femur and tibia possess translucent pores, they are much more difficult to detect in the smaller examples. "In the absence of any experimental evidence, the writer believes that all of this material represents the same species despite the differences in size of the legs and antennae." California Records (Map 65) Kern Co.: Shafter, VII-17-62, on "Timothy" (J. Dean, UCD). Modoc Co.: 8 miles east Tulelake, V-31-61, on "bluegrass," Poa sp. (K. G. Baghott, CDA). Siskiyou Co.: Dunsmuir, VIII-17-65, on Elymus glaucus (T. R. Haig, CDA). Yolo Co.: Davis, VIII-25-58, on "lawn grass"

SPECIES

(H. T. Osborn, CDA); VIII-20-65, on perenne (R. L. Holmes, UCD).

193

Lolium

Map 65. Distribution in California of Heterococcus pulverarius (Newstead).

Marine So» franeitcaSan Matto

HUMOCOCCUS Humococcus Ferris, 1953. Atlas of scale insects of North America, Vol. VI: 370. Genotype. Ripersia hilariae Ferris. Recognition Characters. Pseudococcidae with body form usually broadly oval, as mounted. Dorsum with at least posterior pair of ostioles present. Cerarii usually absent, or with those of anal lobes present, at times represented only by paired, slender setae, in one Nevada species with 5 or 6 pairs. Multilocular disk pores at times present on dorsum. Tubular ducts of the oral-rim type at times absent, at times replaced by ducts of an oral-collar or simple type. Anal ring simplified, with 6 short setae, consisting merely of a narrow, sclerotized ring which in one species bears very small pores; ring in all species borne on dorsum, removed from posterior apex of abdomen by at least its own diameter. Venter with at least a few multilocular disk pores. Tubular ducts of oral-collar and oral-rim type usually present. Circulus absent. Legs normally quite small; claw without denticle except in one species. Antennae 6- or 7-segmented. Notes. The 6- or 7-segmented (usually 6-segmented) antennae, together with the simplified anal ring, presence of only anal lobe cerarii—if present at all—and absence of a circulus are distinctive characteristics of this genus.

OF

FERRIS

The genus Humococcus includes 8 species for North America, 4 of which occur in California.

KEY Species of Humococcus in North America: Adult Females

1.

2(1). 3(2).

4(3).

5(4).

Abdomen with 5 or 6 pairs of cerarii . . ceraricus McKenzie Cerarii absent or with only the anal lobe pair present 2 Anal ring closely surrounded by numerous setae porterae (Cockerell) Anal ring not thus surrounded by setae 3 Oral-rim tubular ducts present on dorsum 4 Oral-rim tubular ducts absent on dorsum . . 6 Dorsal multilocular disk pores present in somewhat irregular band across abdominal segments 8-5 5 Dorsal multilocular disk pores absent across abdominal segments 8-5, at times an occasional such pore near lateral margins on last 2 or 3 abdominal segments 7 At least a few oral-rim and oral-collar

194

6(3).

7(4).

TAXONOMY

OF

SPECIES

tubular ducts on venter of head and thorax; anal ring dorsad from apex of abdomen by 3 or 4 times its own diameter atriplicis Ferris Oral-rim and oral-collar tubular ducts absent ventrally from head' and thorax; anal ring dorsad from apex of abdomen by only slightly more than twice its diameter inornatus McKenzie Oral-collar tubular ducts of the larger size rather numerous, forming a submarginal band on venter from apex of abdomen to head muhlenbergiae Ferris Oral-collar tubular ducts on venter few and confined to a submarginal row on abdomen hilariae (Ferris) Ventral multilocular disk pores present on last 5 or 6 abdominal segments, especially abundant, covering most of area on last 2 segments . . . dasychloae Ferris Ventral multilocular disk pores restricted to last 2 or 3 abdominal segments, comparatively few pores on these segments caritus McKenzie

region and terminal abdominal segments. Body setae short and sparse. Circulus absent. Legs proportionately small; coxa with translucent pores at base; denticle absent on claw. Antennae 7-segmented. Notes. This species is quite closely related to Humococcus caritus McKenzie, but may be distinguished from it by the presence of dorsal multilocular disk pores on abdomen, these structures absent in the species last mentioned. It also resembles H. inornatus McKenzie, but the latter species lacks oral-rim and oral-collar tubular ducts ventrally on head and thorax, these structures present in H. atriplicis. California Records (Map 66) Imperial Co.: East of Holtville, 1-11-66, on Atriplex canescens (R. A. Flock, CDA). Inyo Co.: Death Valley (intercepted in quarantine at Kingman, Arizona) III-1963, on Atriplex hymenelytra (N. O'Connell, CDA); Townes Pass, 1-30-65, on Atriplex sp. (crown) (D. R. Miller, UCD). Kern Co.: Saltdale, IV-26-36, on Atriplex sp. (crown) (G. F. Ferris, UCD).

Humococcus atriplicis Ferris Ferris Atriplex Mealybug (Figure 70) Humococcus atriplicis Ferris, 1953. Atlas of scale insects of North America, Vol. VI:371-372, illus. Type Locality and Host. Near Salt Dale, San Bernardino County, California, on saltbush, Atriplex sp. (Chenopodiaceae). North American Distribution. California, Mexico, and Texas. Additional Hosts. Recorded only on Atriplex spp. External Features and Habitat. According to Ferris (1953), "The only available information indicates that the insects occur on the crowns of the host." Recognition Characters. Adult female, mounted, 1.90 to 2.00 mm long, 1.10 to 1.40 mm wide; body form broadly oval. Dorsum with cerarii absent. Dorsal surface beset with trilocular pores. Multilocular disk pores in a row across abdominal segments 2-8. Oral-rim type tubular ducts small, very few, situated mainly along lateral margins of body. Body setae very small, sparsely distributed. Anal ring quite small, removed by 3 or 4 times its own diameter from posterior apex of abdomen, consisting of a simple, sclerotized ring, interrupted posteriorly, slightly expanded in its lateral areas; bearing 6 exceedingly small setae. Venter with multilocular disk pores situated mostly in rows on all abdominal segments, few scattered on thorax. Trilocular pores and tubular ducts distributed similar to those on dorsum, in addition, a few very small, simple, tubular ducts present especially in head

Humococcus caritus McKenzie McKenzie Salt Grass Mealybug (Figure 71) Humococcus caritus McKenzie, 1960. Hilgardia 29(15):709-711, illus. Type Locality and Host. Lancaster, Los Angeles County, California, on salt grass, Distichlis spicata (Gramineae). North American Distribution. Known only from California. Additional Hosts. None.

TAXONOMY

OF

SPECIES

Fig. 70. Humococcus atriplicis Ferris, collected on saltbush, Atriplex sp. (Chenopodiaceae), near Salt Dale, San Bernardino County, California.

195

196

TAXONOMY

OF

SPECIES

Fig. 71. Humococcus caritus McKenzie, collected on salt grass, Distichlis spicata (Gramineae), at Lancaster, Los Angeles County, California.

TAXONOMY

External Features and Habitat. No available information, although it is assumed that the mealybug occurs on the crown of its host similar to related species. Recognition Characters. Adult female, mounted, 1.50 to 1.90 mm long, 1.00 to 1.10 mm wide; body form elongate oval to slightly pyriform. Dorsum with but one pair cerarii on anal lobes, these represented only by a pair of conical, rather slender, setae, 2-3 slender auxiliary setae, scarcely any concentration of trilocular pores. Trilocular pores very sparse, generally scattered over abdomen, fewer on thorax and head. Multilocular disk pores absent. Small ducts of oral-rim type occur sparingly over dorsal surface. Body setae sparse, all small and slender. Anal ring distinctive for genus, set at about its own diameter from posterior apex of abdomen, consisting merely of a narrow, sclerotized anterior band interrupted along posterior border of opening; bearing 6 small setae, each about as long as diameter of ring. Venter with few multilocular disk pores, confined to posterior portion of body, present on last 2 or 3 abdominal segments. Trilocular pores distributed similar to those on dorsum. Very few, only 3 or 4, small oral-rim tubular ducts situated along body margins. Body setae short and slender, sparse, fewer on thoracic segments than rest of body. Circulus absent. Legs comparatively short; without denticle or tooth on claw. Mouthparts short and broad. Antennae short, 6-segmented. Notes. This species appears to be most closely related to Humococcus atriplicis Ferris, but differs from it principally in the absence of dorsal multilocular disk pores, and in the presence of fewer of these structures ventrally on the abdomen. Califoma Records (Map 67) Los Angeles Co.: Lancaster, VII-28-57, on Distichlis spicata (K. Stevens, CDA).

Map 67. Distribution in California of Humococcus

caritus McKenzie.

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OF

SPECIES

197

Humococcus hilariae (Ferris) Hilaria Mealybug (Figure 72) Ripersia hilariae Ferris, 1919. Stanford Univ. Pubs., Univ. Ser. pp. 32-33, illus. Humococcus hilariae (Ferris), Ferris, 1963. Atlas of scale insects of North America, Vol. VI:373374, illus. Type Locality and Host. On the Jornada del Muerto, 51 miles north of Las Cruces, New Mexico, on grass, Hilaria sp. probably cenchroides (Gramineae). North American Distribution. California, New Mexico, and Texas. Additional Hosts. None to date. External Features and Habitat. According to Ferris (1919), this mealybug was surrounded by a small amount of waxy secretion. The species occurs beneath the sheathing bases of the leaves of its host. Recognition Characters. Adult females, mounted, 1.85 to 2.40 mm long, 0.90 to 1.10 mm wide; body form elongate oval or slightly pyriform. Posterior dorsal ostioles present, anterior pair absent or undetectible in available material. Dorsum with anal lobe cerarii represented merely by a pair of slender setae; remaining cerarii absent. Trilocular pores very few and scattered. Multilocular disk pores very few, present only along submargin of last 2 or 3 abdominal segments, and forward from these along margin to metathorax, absent on remainder of thorax and head. Tubular ducts of oral-collar type very few, situated along margin of last 3 abdominal segments, at times a single one present on head. Body setae very few, all small but increasing somewhat in size and number toward posterior apex of abdomen. Anal ring usually situated somewhat more than its own diameter from posterior apex of abdomen, consisting merely of a narrow, sclerotized band; each of its 6 setae small, scarcely half as long as greatest diameter of ring itself. Venter with multilocular disk pores very few, confined to the area about vulva. Trilocular pores with about same distribution as those on dorsum, somewhat concentrated about spiracles. Tubular ducts of oral-collar type very few, scarcely more than a total of 10, these confined to submarginal row on abdomen, infrequently a single one present on head. Body setae small, fewer than on dorsal side. Circulus absent. Legs proportionately small; coxa bearing numerous small pores; claw without denticle. Antennae 6-segmented. Notes. This species appears somewhat related to Humococcus muhlenbergiae Ferris, a New Mexico species, but differs in presence of at least a few dorsal multilocular disk pores on last 3 abdominal segments, and very few ventral oral-collar type ducts confined to a submarginal row on abdomen. In H. muhlenbergiae, on the other hand, the dorsal multilocular disk pores are absent, and the oral-collar type ducts are

198

TAXONOMY

OF

SPECIES

Fig. 72. Humococcus hilariae (Ferris), on grass, Hilaria rigida (Gramineae), at Seeley, Imperial County, California. (Illustration prepared from type material.)

TAXONOMY

rather numerous, forming an almost continuous submarginal band on venter from posterior apex of abdomen to head. California Records (Map 68) Imperial Co.: Seeley, 111-25-65, on Hilaria rigida (R. A. Flock and L. Pineda, CDA).

Humococcus inornatus McKenzie Inornate Mealybug (Figure 73) Humococcus inornatus McKenzie, 1960. Hilgardia 29(15) :711-713, illus. Humococcus inornatus McKenzie, 1964. Ibid. 35(10):238. Type Locality and Host. Garlic Springs, near Barstow, San Bernardino County, California on undetermined host. North American distribution. To date found only in California. Additional Hosts. A triplex hymenelytra (Chenopodiaceae), Franseria sp. (Compositae), and Haplopappus pinifolius (Compositae), associated with an ant, Camponotus yogi. External Features and Habitat. No information available relative to the external appearance of this mealybug. In 1963 it was intercepted in quarantine at Kingman, Arizona, ' presumably on twigs and branches of A triplex hymenelytra originating from Death Valley, Inyo County, California.

OF

SPECIES

199

Recognition Characters. Adult female, mounted, 1.85 to 3.00 mm long, 1.00 to 1.60 mm wide; body form elongate oval to slightly pyriform. Dorsum with cerarii entirely absent. Trilocular pores rather evenly distributed. Multilocular disk pores in considerable numbers from last to fifth abdominal segments, these for most part extending across segment to lateral margin. A few, small, tubular ducts of oral-rim type situated mostly on abdomen, very few located on head, sternal region, and along thoracic margin. Oral-collar tubular ducts sparsely situated on last abdominal segment. Dorsal surface with but very few body setae, these small and slender. Anal ring quite small, removed by about 2Vi times its own diameter from posterior apex of abdomen, consisting of a simple, sclerotized ring, interrupted posteriorly, slightly expanded in lateral areas; surrounded by 6 small setae, each less than one-half diameter of ring itself. Venter with multilocular disk pores in considerable numbers from posterior apex to third abdominal segment, these mostly in rows extending across segments to lateral margin, fewer on segments 3 and 4. Trilocular pores distributed similar to those on dorsum. Oral-rim tubular ducts similar in size to those on dorsum, situated especially on terminal segments of abdomen, none present in sternal areas or on head. A few, small, simple, tubular ducts of oral-collar type sparsely situated on eighth to fifth abdominal segments. Body setae short, slender, sparse, fewer on thorax than rest of body. Circulus absent. Legs comparatively short; claw with small denticle near tip, sometimes obscure. Mouthparts short and relatively broad. Antennae 7-segmented. Notes. This species is rather closely related to Humococcus atriplicis Ferris, but differs in lacking oral-rim and oral-collar tubular ducts ventrally on head and thorax; and anal ring surrounded by, but not bearing, 6 small setae. On the other hand, H. atriplicis, possesses ventral oral-rim and oral-collar tubular ducts; and the anal ring bears 6 small setae. The construction of the anal ring of H. inornatus does not seem to match any other species included in this group. California Records (Map 69) Inyo Co.: Death Valley (intercepted in quarantine at Kingman, Arizona), III—1963, on A triplex hymenelytra (N. O'Connell, CDA). San Bernardino Co.: 5 miles north Baker, IV-1363, on Franseria sp. (D. R. Miller, UCD); Garlic Springs, near Barstow, IV-8-35, on ? (J. D. Maple, UCD). San Diego.: 12 miles east Borrego Springs, X I I 24-46, on Franseria dumosa (D. R. and J. F. Miller, UCD).

200

TAXONOMY

OF

SPECIES

Fig. 73. Humococcus inornatus McKenzie, collected on Franseria sp. (Compositae), north of Baker, San Bernardino County, California. (Illustration prepared from type material.)

TAXONOMY

Sitkitjou

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SPECIES

201

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HYPOGEOCOCCUS Hypogeococcus Rau, 1938. Canad. Ent. 7 0 ( 8 ) : 157165, illus. Genotype. Hypogeococcus barbarae Rau. Recognition Characters. Pseudococcidae with body form variable, from narrowly oval to rather strongly ovoid, as mounted. Dorsal ostioles present. Cerarii confined to last 5 or 6 abdominal segments. Multilocular disk pores present on dorsum and venter. Tubular ducts of both surfaces simple, with oral collar, at most, only slightly developed. Dorsum of abdomen beset with stout setae which are nearly or quite as large as those of cerarii. Anal ring situated at posterior apex of body, with no special distinctive features. Circulus present, circular, apparently entirely enclosed within boundaries of fourth abdominal segment. Legs well developed; tarsal claw without denticle or tooth on plantar surface. Antennae 7-segmented. Notes. In the character of the enlarged, stout dorsal body setae, this genus appears distantly related to Nipaecoccus Sulc and Amonostherium Morrison and Morrison. Two species of Hypogeococcus are known from North America, 1 of which occurs in California. It is possible that these 2 species should not be considered as congeneric, yet, as here interpreted, they are sufficiently similar to warrant this assignment.

RAU

KEY Species of Hypogeococcus in North America: Adult Females (Slightly modified from Ferris, 1953) Cerarian conical setae definitely differentiated from enlarged, dorsal, body setae on abdomen barbarae Rau Cerarian conical setae only in part differentiated from enlarged, dorsal, body setae on abdomen spinosus Ferris

Hypogeococcus spinosus Ferris Spinose Mealybug (Figure 74) Hypogeococcus spinosus Ferris, 1953. Atlas of scale insects of North America, Vol. VI: 382-383, illus. Type Locality and Host. Delhi, Merced County, California, on undetermined cactus (Cactaceae), presumably growing under cultivation. North American Distribution. California and Texas. Additional Hosts. Unidentified "cactus" and Echinocereus sp. (Cactaceae). External Features and Habitat. Specimens inter-

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Fig. 74. Hypogeococcus spinosus Ferris, collected on undetermined "cactus" (Cactaceae), at Delhi, Merced County, California.

TAXONOMY

cepted in quarantine on cactus from Texas were collected on the stem of the host. No other information available at this time. Recognition Characters. Adult female, mounted, 1.40 to 1.50 mm long, 1.10 to 1.20 mm wide; body form strongly ovoid. Cerarii scarcely differentiated from stout setae of dorsum, with which they are more or less confluent, except on anal lobes. Anal lobe cerarius with 3 or 4 stout, conical setae. Trilocular pores apparently entirely absent. Multilocular pores present over entire dorsum, arranged in irregular transverse row or rows across anterior and posterior margins of abdominal segments, scattered on thorax and head. Short tubular ducts with a small oral-collar present in bands across abdominal segments, quite sparingly scattered over thoracic and head areas. Enlarged body setae of abdomen extending in a narrow band across body to extreme lateral margins, present on third to eighth segments, mingled with few quite long, slender setae. Anterior to abdomen, setae quite small, slender, scattered. Anal ring almost quadrilateral, strongly sclerotized, with numerous small pores; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores situated rather sparsely throughout. Trilocular pores absent. Tubular ducts similar to those on dorsum, few, scattered in all areas. Body setae small, slender, sparsely distributed throughout. Circulus circular, moderately large. Legs rather slender; denticle absent on claw. Antennae 7-segmented.

OF

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203

It appears related to Hypogeococcus barbarae Rau, a New York species, but differs in having abdominal cerarian setae less differentiated from the enlarged dorsal body setae, whereas in barbarae the abdominal cerarian setae are markedly differentiated from the more slender dorsal body setae. California Records (Map 70) Merced Co.: Delhi (in nursery), IV-6-51, on "cactus" (J. J. Kalstrom and E. Burton, CDA). San Diego Co.: Carlsbad, IX-16-59, on Echinocereus sp. (H. T. Oldham, CDA); San Diego, I I 16-57, on "cactus" (D. Sprague and G. W. Schwegel, CDA).

Map 70. Distribution in California of Hypogeococcus

Ferris.

spionosus

San HaneiiccSan Matto

Notes. Some of the taxonomic features which distinguish this species from other California mealybugs is the presence of numerous stout, conical, setae on dorsal surface of abdomen which are approximately the same size and shape as those of the cerarii, and with 3 or 4 of these structures on each anal lobe cerarius.

MISERICOCCUS FERRIS Misericoccus Ferris, 1953. Atlas of scale insects of North America, Vol. VI: 384. Genotype. Ripersia arenaria Doane and Steinweden. Recognition Characters. Pseudococcidae with body form elongate oval. Dorsal ostioles represented only by the posterior pair, anterior pair absent or not detectible. Cerarii represented only by anal lobe pair, these consisting merely of paired, small, conical setae. Trilocular pores absent, replaced by multilocular disk pores over entire surface. Tubular ducts absent. Anal ring reduced, with few pores, its sclerotization incomplete, ring open anteriorly for about one-third of its extent; its 6 setae small, shorter than diameter of ring itself.

Venter with trilocular pores absent, replaced by multilocular disk pores over whole surface. Tubular ducts very few, confined to posterior abdominal segments, very few on head. Circulus absent. Legs very small; claw without denticle or tooth on plantar surface. Antennae 6-segmented. Notes. Ferris (1953) states the following about this genus: "The author dislikes the naming of a new genus for this, but not to do so is unduly to complicate the definition of other genera. This genus is perhaps closest to Humococcus but to refer it to this genus produces what seems to be an otherwise quite natural group."

TAXONOMY

cepted in quarantine on cactus from Texas were collected on the stem of the host. No other information available at this time. Recognition Characters. Adult female, mounted, 1.40 to 1.50 mm long, 1.10 to 1.20 mm wide; body form strongly ovoid. Cerarii scarcely differentiated from stout setae of dorsum, with which they are more or less confluent, except on anal lobes. Anal lobe cerarius with 3 or 4 stout, conical setae. Trilocular pores apparently entirely absent. Multilocular pores present over entire dorsum, arranged in irregular transverse row or rows across anterior and posterior margins of abdominal segments, scattered on thorax and head. Short tubular ducts with a small oral-collar present in bands across abdominal segments, quite sparingly scattered over thoracic and head areas. Enlarged body setae of abdomen extending in a narrow band across body to extreme lateral margins, present on third to eighth segments, mingled with few quite long, slender setae. Anterior to abdomen, setae quite small, slender, scattered. Anal ring almost quadrilateral, strongly sclerotized, with numerous small pores; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores situated rather sparsely throughout. Trilocular pores absent. Tubular ducts similar to those on dorsum, few, scattered in all areas. Body setae small, slender, sparsely distributed throughout. Circulus circular, moderately large. Legs rather slender; denticle absent on claw. Antennae 7-segmented.

OF

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It appears related to Hypogeococcus barbarae Rau, a New York species, but differs in having abdominal cerarian setae less differentiated from the enlarged dorsal body setae, whereas in barbarae the abdominal cerarian setae are markedly differentiated from the more slender dorsal body setae. California Records (Map 70) Merced Co.: Delhi (in nursery), IV-6-51, on "cactus" (J. J. Kalstrom and E. Burton, CDA). San Diego Co.: Carlsbad, IX-16-59, on Echinocereus sp. (H. T. Oldham, CDA); San Diego, I I 16-57, on "cactus" (D. Sprague and G. W. Schwegel, CDA).

Map 70. Distribution in California of Hypogeococcus

Ferris.

spionosus

San HaneiiccSan Matto

Notes. Some of the taxonomic features which distinguish this species from other California mealybugs is the presence of numerous stout, conical, setae on dorsal surface of abdomen which are approximately the same size and shape as those of the cerarii, and with 3 or 4 of these structures on each anal lobe cerarius.

MISERICOCCUS FERRIS Misericoccus Ferris, 1953. Atlas of scale insects of North America, Vol. VI: 384. Genotype. Ripersia arenaria Doane and Steinweden. Recognition Characters. Pseudococcidae with body form elongate oval. Dorsal ostioles represented only by the posterior pair, anterior pair absent or not detectible. Cerarii represented only by anal lobe pair, these consisting merely of paired, small, conical setae. Trilocular pores absent, replaced by multilocular disk pores over entire surface. Tubular ducts absent. Anal ring reduced, with few pores, its sclerotization incomplete, ring open anteriorly for about one-third of its extent; its 6 setae small, shorter than diameter of ring itself.

Venter with trilocular pores absent, replaced by multilocular disk pores over whole surface. Tubular ducts very few, confined to posterior abdominal segments, very few on head. Circulus absent. Legs very small; claw without denticle or tooth on plantar surface. Antennae 6-segmented. Notes. Ferris (1953) states the following about this genus: "The author dislikes the naming of a new genus for this, but not to do so is unduly to complicate the definition of other genera. This genus is perhaps closest to Humococcus but to refer it to this genus produces what seems to be an otherwise quite natural group."

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Misericoccus arenarius (Doane and Steinweden) Pear-Shaped Mealybug (Figure 75) Ripersia arenaria Doane and Steinweden, 1927. PanPacific Ent. 4 ( 2 ) : 85-86, illus. Misericoccus arenarius (Doane and Steinweden), Ferris, 1953. Atlas of scale insects of North America, Vol. VI: 384-385, illus. Type Locality and Host. Near Salt Lake City, Sa't Lake County, Utah, on Bromus arenarius (Gramineae). North American Distribution. California and Utah. Additional Hosts. Bromus rubens, Bromus tectorum, "grass," Hordeum leporinum, Hordeum sp., Lolium sp., and Stipa sp. (all Gramineae). External Features and Habitat. According to the original description, the adult females of this mealybug are covered with a considerable amount of white, cottony secretion. It occurs on the roots of its host. It has also been found under rocks and in one instance was attended by the ant, Formica camponoticeps. Recognition Characters. Adult female, mounted, 1.40 to 2.60 mm long, 1.00 to 1.40 mm wide; body form elongate oval. Dorsum with cerarii represented only by anal lobe pair, these consisting merely of paired, small, conical setae. Trilocular pores absent, replaced by multilocular disk pores situated over entire dorsal body surface. Minute circular pores scattered. Tubular ducts of any kind absent. Body setae extremely small, relatively few, becoming slightly larger and more or less lanceolate on abdomen. Anal ring situated at posterior apex of abdomen, sclerotization quite broad but enveloping only posterior two-thirds, anterior portion open, pores small and oval; its 6 setae scarcely more than half as long as diameter of ring. Venter with trilocular pores absent, replaced by multilocular disk pores over whole surface. Minute circular pores scattered. Tubular ducts very few, confined to last 4 or 5 abdominal segments, very few on head. Body setae similar to those on dorsum, some slightly longer. Circulus absent. Legs very small; denticle or tooth on claw absent. Antennae 6-segmented. Notes. This monotypic genus is quite different from most other California mealybugs in the absence of trilocular pores and in the presence of a 6-segmented antenna. It shares with certain California members of Heterococcus the absence of the trilocular pores, but differs in having only multilocular disk pores present

instead of both multilocular and quinquelocular pores, which characterizes the genus last mentioned. California Records (Map 71) Fresno Co.: Fresno, IV-17-56, on Lolium sp. (G. G. Beevor and H. Bronson, CDA). Los Angeles Co.: Lancaster, IV-6-64, on "grass" (roots) (G. G. Beevor and J. L. Johnson, CDA); Neenach, Lancaster, IV-6-61, on Hordeum leporinum (roots) (G. G. Beevor, CDA). Modoc Co.: 6 miles east Alturas, VI-27-63, on Bromus tectorum (roots and under rocks) (D. R. Miller, UCD); near Cedarville, V-21-62, "under rock attended by ant, Formica camponoticeps," (T. R. Haig and R. F. Wilkey, CDA). Nevada Co.: Boca Dam, VI-26-64, on Bromus sp. (roots) (D. R. Miller, UCD). San Benito Co.: Pacheco Pass, III—22—15, on Stipa sp. (roots) (G. F. Ferris, UCD). San Diego Co.: 0.5 mile north Oceanside, 1-29-64, on Bromus rubens (roots and crowns) (D. R. Miller, UCD). San Luis Obispo Co.: Carrizo Plain, III—30—35, on Hordeum sp. (roots) (G. F. Ferris, UCD).

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Fig. 75. Misericoccus arenarius (Doane and Steinweden), collected on grass, Bromus rubens (Gramineae), north of Oceanside, San Diego County, California. (Illustration prepared from type material.)

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NIPAECOCCUS Nipaecoccus Sulc, 1945. Acta Soc. Sci. Nat. Morav. 1 7 ( 3 ) : 1-48, illus. Nipaecoccus Sulc, Ferris, 1950. Atlas of scale insects of North America, Ser. V: 103. Nipaecoccus Sulc, Williams, 196-2. Bull. Brit. Mus. (Nat. Hist.) Ent. 12(1) :31. Genotype. Dactylopius nipae Maskell. Recognition Characters. Pseudococcidae with body form, as mounted, usually quite rotund. Dorsal ostioles tend to be more or less reduced, anterior pair at times seemingly absent, posterior pair at times being a mere slit that is difficult to recognize. Cerarii reduced in number; anal lobe pair always with but 2 large, conical, or somewhat lanceolate setae, others also with 2 setae of same type but these often rather widely separated, becoming more so toward thorax, finally becoming unrecognizable and disappearing entirely from anterior portion of body. Dorsal multilocular disk pores and tubular ducts present or absent. Body setae on dorsum always, at least in part, of same shape and as large as, or only slightly smaller than, those of cerarii. Anal ring of normal form. Circulus present or absent. Multilocular disk pores always present, at least about vulva. Tubular ducts always on venter, variously disposed. Legs normally quite small and short. Antennae normally 7-segmented. Notes. This genus seems rather closely related to Amonostherium Morrison and Morrison; however, the type of Amonostherium lacks definite cerarii other than those of the anal lobes, and possesses a large number of setae in the anal lobe cerarii. Nipaecoccus, on the other hand, has several cerarii, and the anal lobe cerarii possesses not more than 2 setae. The differences between the two genera are distinct and can be adequately expressed. Species of Nipaecoccus in North America are widely spread. The species occur for the most part in tropical and subtropical areas around the world. The genus Nipaecoccus includes 5 species for North America, 2 of which occur in California.

KEY Species of Nipaecoccus in North America: Adult Females

1. 2(1).

3(1).

(Modified from Ferris, 1953) Circulus present 3 Circulus absent 2 Dorsum entirely without tubular ducts and multilocular disk pores ledi (Cockerell) Dorsum with both tubular ducts and multilocular disk pores filamentosus (Cockerell) Dorsum with a distinct, submarginal cluster of conical setae near base of each antenna nipae (Maskell)

SULC

4(3).

Dorsum without such cephalic clusters of conical setae 4 Dorsum with extensive bare areas which contain neither pores nor setae; apparently tubular ducts absent on dorsum aurilanatus (Maskell) Dorsum without such bare areas, and with considerable numbers of small, tubular ducts vastator (Maskell)

Nipaecoccus aurilanatus (Maskell) Golden Mealybug (Figure 76; Color Plate III) Dactylopius aurilanatus Maskell, 1889. Trans. New Zeal. Inst. 22:151-152, illus. Dactylopius aurilanatus Maskell, Craw, 1896. Calif. State Bd. Hort., Bien. Rpt. 5 (1895-96), p. 45, illus. Dactylopius aurilanatus Maskell, 1898. Trans. New Zeal. Inst. 30(1897) :320. Dactylopius aurilanatus Maskell, Froggat, 1897. Dept. Agr. N.S. Wales Misc. Pub. 175:4-5. Dactylopius aurilanatus Maskell, Fuller, 1901. Natal Dept. Agr., First Rpt. Ent., p. 106. Pseudococcus aurilanatus (Maskell), Essig, 1914. Calif. State Comn. Hort. Monthly Bull. 3 ( 3 ) : 108-109, illus. Pseudococcus aurilanatus (Maskell), Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., p. 38, illus. Nipaecoccus aurilanatus (Maskell), Ferris, 1950. Atlas of scale insects of North America, Ser. V: 104-105, illus. Type Locality and Host. Auckland, New Zealand, on Bunya-Bunya, Araucaria bidwillii (Araucariaceae). North American Distribution. Known only from California. Additional Hosts. Agathis sp. (Araucariaceae), Araucaria excelsa, Araucaria spp. (Araucariaceae), and Diplacus longiflorus (Scrophulariaceae). External Features and Habitat. Maskell's original description of the external features of this pseudococcid are quite clear and may be quoted as follows: "Adult female slightly elongated, nearly globular; of a rich, dark-purple colour, bearing on the dorsum a longitudinal band of bright golden coloured meal, with small patches of similar meal often visible at the edges. . . . The eggs, which are also purple, are laid in a mass behind the insect, in a thin white cottony web, the mass thus having a general dark grey appearance" (see col. pi. III). This mealybug occurs on the small branches and needles of its host. Recognition Characters. Adult female, mounted, 2.00 to 3.00 mm long, 1.20 to 2.00 mm wide; body form quite rotund. Dorsum with not more than 6 recognizable pairs

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Fig. 76. Nipaecoccus aurilanatus (Maskell), collected on Norfolk Island pine, Araucaria excelsa (Araucariaceae), at Santa Ana, Orange County, California. (Illustration prepared from specimens collected on Araucaria bidwilli [Araucariaceae], on campus at Stanford University, Santa Clara County, California.)

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of cerarii including those on anal lobes. Anal lobe cerarius with 2 rather small, slender, conical setae, few trilocular pores, surrounded by small, rather indistinct sclerotized area. Anterior to anal lobes not more than five pairs of cerarii can be recognized; in these the setae are smaller than those of anal lobes and are somewhat separated, especially in anteriormost pairs. Dorsal ostioles quite small, indistinct. Trilocular pores scattered on dorsum. Minute circular pores scattered. Tubular ducts absent. Dorsal body setae for most part same shape as those of lateral cerarii, some of these on posterior abdominal segments about as large as setae of anal lobes, usually quite few, occur mostly along midline in very indefinite submarginal clusters. Greater part of dorsum destitute of either trilocular pores or setae, only head region having both in abundance—the areas of pores and setae evidently correspond to dorsal patches of yellow wax. Anal ring situated at posterior apex of abdomen; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores quite numerous from vulva forward to circulus, situated mostly in midregion. Trilocular pores very few. Minute circular pores scattered. A very few, small, oral-collar type ducts situated throughout median areas and along lateral areas of abdominal segments. Body setae small and slender, sparsely distributed. Circulus present. Legs small but slender; tibiotarsal portions equalling in length trochanter-femoral portion. Antennae 7- or 8-segmented. Notes. This is the "golden mealybug," which is a serious pest of Araucaria grown as ornamentals in California. The species is rather closely related to N. nipae (Maskell), the only other known segregate of this genus in California, but is easily differentiated by the presence of numerous ventral multiloculars on all abdominal segments; the absence of a dorsal cephalic cluster of conical setae above antennal base; and a coniferous host. Nipaecoccus nipae, on the other hand, possesses comparatively few ventral multiloculars from apical to posterior margin of sixth abdominal segments; an obvious dorsal cephalic cluster of conical setae above base of each antenna; and many hosts, especially palms, and is rarely, if ever, on conifers. California Records (Map 72) California, date ?, host ?, (E. M. Ehrhorn, USNM). Los Angeles Co.: Beverly Hills, 1-7-58, on Araucaria excelsa (D. Ferrell, CDA); Pasadena, X I I 8-36, on Araucaria sp. (H. J. Quayle, CDA); date ?, on Araucaria sp. (A. D. Borden, UCD); San Gabriel, VII-29-36, on Araucaria sp. (L. E. Myers, CDA); Whittier, V-1912, on Diplacus longiflorus (P. H. Timberlake, USNM). Orange Co.: Santa Ana, X - 8 - 1 2 , on Araucaria excelsa (R. K. Bishop, USNM). San Mateo Co.: Menlo Park, VIII-24-62, on Agathis sp. (E. L. Labadie, CDA). Santa Clara Co.: Mountain View, IX-15-16, on Araucaria sp. (W. L. Garlieph, UCD); San Jose,

VII-22-42, on Araucaria bidwilli (M. R. Bell, UCD); Stanford University Campus, VI-1945, on Araucaria bidwilli (G. F. Ferris, UCD); II-8-27, on Araucaria sp. (L. E. Myers, L E M ) ; 1937, on Araucaria sp. (F. S. Stickney, USNM).

Nipaecoccus nipae (Maskell) Coconut Mealybug (Figure 77; Color Plate II) Dactylopius nipae Maskell, 1892. Trans. New Zeal. Inst. 25:232, illus. Dactylopius pseudonipae Cockerell, 1897. Sei. Gossip (n.s.) 3:189,302. Ripersia serrata Tinsley, 1900. Canad. Ent. 32(3)66. Dactylopius dubia Maxwell-Lefroy, 1903. West Indies Imp. Dept. Agr. (Pam. Ser.) 22:26, illus. Pseudococcus nipae (Maskell) Fernald, 1903. Mass. Agr. Expt. Sta. Spec. Bull. 88, p. 107. Trechocorys nipae (Maskell), Kirkaldy, 1904. Hawaii. For. and Agr. 1 ( 6 ) : 154. Pseudococcus nipae (Maskell), Marchal, 1908. Ann. Soc. Ent. de France 77(8):236-239, illus. Pseudococcus pseudonipae (Cockerell), Essig, 1914. Calif. State Comn. Hort. Monthly Bull. 3:120121, illus. Pseudococcus nipae (Maskell), Green, 1917. Ent. Mon. Mag. 53:262-263, illus. Pseudococcus nipae (Maskell), Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., pp. 49-50, illus. Nipaecoccus nipae (Maskell), Sulc, 1945. Acta Soc. Sei. Nat. Morav 1 7 ( 3 ) : 1-48, illus.

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Fig. 77. Nipaecoccus tiipae (Maskell), collected on palm, Kentia sp. (Palmaceae), at San Francisco (Golden Gate Park), San Francisco County, California. (Illustration prepared from specimens collected on Kentia sp. [Palmaceae], at Marysville [nursery], Yuba County, California.)

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Pseudococcus nipae (Maskell), Zimmerman, 1948. Insects of Hawaii, Univ. Hawaii Press, 5 : 2 2 9 235, illus. Nipaecoccus nipae (Maskell), Ferris, 1950. Atlas of scale insects of North America, Ser. V: 109-111, illus. Nipaecoccus nipae (Maskell), Ferris, 1953. Ibid., Vol. VI: 391. Nipaecoccus nipae (Maskell), Williams, 1962. Bull. Brit. Mus. (Nat. Hist.) Ent. 1 2 ( 1 ) : 3 1 . Type Locality and Host. Demerara, British Guiana, on an aquatic palm, Nipa fruticans (Palmaceae). North American Distribution. California, Florida, Hawaii, Louisiana, Mexico, and Puerto Rico. Additional Hosts. This mealybug has been recorded especially from palms, most commonly of the genus Kentia, in greenhouses in Northern latitudes in both Europe and North America. A few other recorded hosts include: Annona sp. (Annonaceae), Asparagus sp. (Liliaceae), Artocarpus altilis (Moraceae), Canna sp. (Cannaceae), Coccoloba sp. (Polygonaceae), Dracaena sp. (Agavaceae) Ficus benghalensis (Moraceae), Monstera sp. (Araceae), Morus sp. (Moraceae), Persea sp. (Lauraceae), Psidium sp. (Myrtaceae), Sterculia urens (Sterculiaceae), Straussia sp. (Rubiaceae), Strelitzia sp. (Musaceae), Urera sp. (Urticaceae), and Vitis sp. (Vitaceae). External Features and Habitat. Maskell's original description of this species includes a good account of its external features. His remarks are as follows: "Adult female dark-red in colour, flattish, subcircular; diameter about 1 / 3 0 in. exclusive of the cotton; excreting on the dorsum whitish or yellowish cottony meal, and all round the margin subcylindrical tassels of yellowish cotton, which are sometimes equally long all round, sometimes longer posteriorly; the dorsal meal is often arranged in rows in little granular masses" (see col. pi. I I ) . This mealybug infests the foliage of its host. Recognition characters. Adult female, mounted, 1.90 to 2.20 mm long, 1.20 to 2.00 mm wide; body form normally rotund; derm at maturity frequently quite strongly pigmented. Dorsum with definite cerarii confined to abdominal segments. Anal lobe cerarius with 2 moderately large, conical setae, few trilocular pores, these borne in quite distinct sclerotization. Abdominal cerarii anterior to anal lobes with cerarian setae slightly smaller, retaining conical form, these usually rather widely separated, thoracic areas and head without definite cerarii, although some irregular clustering of setae noticeable along margin, cluster on head at times assuming at most a slight appearance of being cerarii. Dorsal ostioles well developed. Dorsum with trilocular pores rather numerous, evenly distributed. Minute circular pores scattered. Tubular ducts absent. Dorsal body setae almost exclusively of same form as those of cerarii, tending to be smaller and arranged in 4 longitudinal rows. It seems logical to assume that the dorsal clumps of wax are formed from trilocular pores in connection with

rows of conical setae. Anal ring somewhat removed from posterior apex of abdomen; its 6 setae about twice as long as diameter of ring. Venter with very few multilocular disk pores, these situated between vulva and circulus. Entire venter very sparsely provided with trilocular pores. Minute circular pores scattered. Very few small tubular ducts scattered on abdomen and sternal areas. Body setae small, slender, sparsely distributed. Circulus present, small and circular. Legs small but not especially stout; hind coxa with a few translucent pores at base. Antennae usually 7-segmented, rarely with 6 segments. Notes. According to Ferris (1950), "The conclusions concerning the identity of this species are not especially satisfactory, and the identification and synonomy are based upon circumstantial evidence and a theory of probabilities. The original description of nipae as given by Maskell applies, as far as it goes, very definitely to the species which occurs commonly on palms in greenhouses in the United States. There is no indication of the presence of a second species." Ferris further remarked: "The situation in regard to Dactylopius pseudonipae Cockerell is equally unsatisfactory. Dr. Harold Morrison, who has very kindly looked into the matter, offers the following notes. The description of pseudonipae is in Science Gossip (New Series), III (1897), p. 302. Cockerell there states that the specimens upon which he based this name were sent to him by Alexander Craw, from a California greenhouse, but these specimens do not exist in the National Collection and specimens from another source have there been erroneously indicated as the types. So the status of the species is quite obscure." There has been some suggestion that possibly two species are involved in material identified as nipae— one with waxy secretions which are white, the other yellowish. Maskell noted that both colors were evident in his material. In recent conversation with J. W. Beardsley, who has done excellent taxonomic work on pseudococcid males, I was informed that there are distinct differences in the males of this socalled "nipae" complex. Nipaecoccus nipae differs from N. aurilanatus in having fewer ventral multiloculars, these confined to the last 4 abdominal segments; an obvious dorsal cephalic cluster of conical setae above base of each antenna; and many hosts, especially palms, rarely if ever found on conifers. N. aurilanatus, on the other hand, has numerous ventral multiloculars on all abdominal segments; absence of a dorsal cephalic cluster of conical setae above antennal base; and a coniferous host. California Records (Map 73) California (quarantined at New York), IV-1-35, on "palm" (C. G. Lathom, USNM); California, (quarantined at Rio Piedras, P. R . ) , IV-5-35, on ?, (G. N. Wolcott, USNM). Alameda Co.: Oakland, 11-13-17, on "palm" (E. O. Essig, CAS); 11-26-30, on Kentia sp. F. J. March, CDA).

TAXONOMY Kern Co. : Bakersfield, 111-29-40, on Kentia sp. C. S. Morley, CDA) ; III-9-39, on Philodendron sp (C. S. Morley, CDA). Los Angeles Co. : Los Angeles, VII-7-26, on Kentia sp. (L. E. Myers, LEM); Montebello, 1-19-35, on Kentia sp. (J. D. Maple, UCD); Montebello, 11-29-36, on Kentia sp. (R. Wilcox, CDA) ; Pasadena (Huntington Estate), IV-1918, on ? (R. W. Doane, UCD.) Orange Co.: Olive, X-17-38, on "palm" (R. J. Bumgardner, CDA). San Bernardino Co.: Ontario, IV-2-31, on Phoenix sp. (G. A. Pohl, CDA); Upland, XI-15-41, on Strelitzia sp. (C. Hayward, CDA). San Diego Co.: La Jolla (quarantined at San Ysidro), VI-17-36, on Livistona sp. (Schoening and Chatterly, USNM); V-27-31, on "palm" (Howard and Schoening, USNM); La Mesa, VI-24-40, on Kentia sp. (R. R. McLean, CDA); Lemon Grove, IV-1-57, on ? (K. Sims and G. W. Schwegel, CDA); San Diego, II-2-60, on Livistona sp. (S. E. Flanders, UCD); X-ll-63, on "palm" (C. R. Caballero, CDA); San Diego (Coronado Island), III-8-07, on "palm" (P. C. Ting, CDA); San Diego, 111-26-40, on Phoenix canadensis (J. W. Dixon, CDA). San Francisco Co.: San Francisco (Golden Gate Park), VI-10-37, on Kentia sp. (P. C. Ting, CDA); IX-28-44, on "palm" (M. R. Bell, CDA). Santa Barbara Co.: Santa Barbara, IX-3-11, on Kentia sp. ?, (Collector ?, UCD); XI-1924, on "palm" (F. C. Greer, UCD); VIII-11-58, on Kentia sp. (M. Suskin, CDA); VIII-15-61, on Kentia sp. (M. Suskin, CDA) ; XII-28-62, on Cordyline .yp. (M. Suskin, CDA).

SPECIES

211

Ventura Co.: Camarillo, III-1-47, on Kentia sp. (P. B. Travis, CDA). Yuba Co. : Marysvills, X-28-44, on Kentia sp. (A. W. Worledge, CDA).

Map 73. Distribution in California of Nipaecoccus nipae (Maskell).

Marin-* San Franei*eoSanMatto Santa Cruz

PALUDICOCCUS Paludicoccus Ferris, 1953. Atlas of scale insects of North America, Vol. VI: 392. Genotype. Sphaerococcus disticlium Kuwana. Recognition Characters. Pseudococcidae included in family only by presence of trilocular pores, tubular ducts which are of the type that occurs on other forms that seem definitely to be pseudococcid, and character of antennae of first-stage nymph. Dorsal ostioles absent in any stage. Cerarii absent. Trilocular pores present on dorsum. Multilocular disk pores present both dorsally and ventrally. Tubular ducts present on both body surfaces in broad zone around margin of body, these ducts closed at inner extremity by strongly sclerotized bar. Anal ring situated on dorsum, removed by more than its own diameter from posterior apex of abdomen, strongly sclerotic, with very few small pores; its 6 setae barely half as long as diameter of ring. Venter with trilocular pores very sparsely distributed throughout. Spiracles with large, strongly sclerotic apodeme, without a sclerotized peritreme.

OF

FERRIS

Circulus absent. Legs absent at maturity and probably in penultimate stage. Antennae at maturity reduced to mere 3-segmented stubs. First-stage nymph with 6-segmented antennae, which are entirely of pseudococcid type; with trilocular pores; without dorsal ostioles. Notes. This is one of the more extreme pseudococcid types. Even though it lacks dorsal ostioles in any of the developmental stages, the presence of trilocular pores and tubular ducts of a type characteristic of the family make its assignment here seem entirely justified.

Paludicoccus distichlium (Kuwana) Kuwana Brown Mealybug (Figure 78) Pseudolecanium disticlium Kuwana, 1902. Ent. News 13:134-135, illus. Sphaerococcus disticlium (Kuwana), Fernald, 1903. Mass. Agr. Expt. Sta. Spec. Bull. 88, p.85.

TAXONOMY Kern Co. : Bakersfield, 111-29-40, on Kentia sp. C. S. Morley, CDA) ; III-9-39, on Philodendron sp (C. S. Morley, CDA). Los Angeles Co. : Los Angeles, VII-7-26, on Kentia sp. (L. E. Myers, LEM); Montebello, 1-19-35, on Kentia sp. (J. D. Maple, UCD); Montebello, 11-29-36, on Kentia sp. (R. Wilcox, CDA) ; Pasadena (Huntington Estate), IV-1918, on ? (R. W. Doane, UCD.) Orange Co.: Olive, X-17-38, on "palm" (R. J. Bumgardner, CDA). San Bernardino Co.: Ontario, IV-2-31, on Phoenix sp. (G. A. Pohl, CDA); Upland, XI-15-41, on Strelitzia sp. (C. Hayward, CDA). San Diego Co.: La Jolla (quarantined at San Ysidro), VI-17-36, on Livistona sp. (Schoening and Chatterly, USNM); V-27-31, on "palm" (Howard and Schoening, USNM); La Mesa, VI-24-40, on Kentia sp. (R. R. McLean, CDA); Lemon Grove, IV-1-57, on ? (K. Sims and G. W. Schwegel, CDA); San Diego, II-2-60, on Livistona sp. (S. E. Flanders, UCD); X-ll-63, on "palm" (C. R. Caballero, CDA); San Diego (Coronado Island), III-8-07, on "palm" (P. C. Ting, CDA); San Diego, 111-26-40, on Phoenix canadensis (J. W. Dixon, CDA). San Francisco Co.: San Francisco (Golden Gate Park), VI-10-37, on Kentia sp. (P. C. Ting, CDA); IX-28-44, on "palm" (M. R. Bell, CDA). Santa Barbara Co.: Santa Barbara, IX-3-11, on Kentia sp. ?, (Collector ?, UCD); XI-1924, on "palm" (F. C. Greer, UCD); VIII-11-58, on Kentia sp. (M. Suskin, CDA); VIII-15-61, on Kentia sp. (M. Suskin, CDA) ; XII-28-62, on Cordyline .yp. (M. Suskin, CDA).

SPECIES

211

Ventura Co.: Camarillo, III-1-47, on Kentia sp. (P. B. Travis, CDA). Yuba Co. : Marysvills, X-28-44, on Kentia sp. (A. W. Worledge, CDA).

Map 73. Distribution in California of Nipaecoccus nipae (Maskell).

Marin-* San Franei*eoSanMatto Santa Cruz

PALUDICOCCUS Paludicoccus Ferris, 1953. Atlas of scale insects of North America, Vol. VI: 392. Genotype. Sphaerococcus disticlium Kuwana. Recognition Characters. Pseudococcidae included in family only by presence of trilocular pores, tubular ducts which are of the type that occurs on other forms that seem definitely to be pseudococcid, and character of antennae of first-stage nymph. Dorsal ostioles absent in any stage. Cerarii absent. Trilocular pores present on dorsum. Multilocular disk pores present both dorsally and ventrally. Tubular ducts present on both body surfaces in broad zone around margin of body, these ducts closed at inner extremity by strongly sclerotized bar. Anal ring situated on dorsum, removed by more than its own diameter from posterior apex of abdomen, strongly sclerotic, with very few small pores; its 6 setae barely half as long as diameter of ring. Venter with trilocular pores very sparsely distributed throughout. Spiracles with large, strongly sclerotic apodeme, without a sclerotized peritreme.

OF

FERRIS

Circulus absent. Legs absent at maturity and probably in penultimate stage. Antennae at maturity reduced to mere 3-segmented stubs. First-stage nymph with 6-segmented antennae, which are entirely of pseudococcid type; with trilocular pores; without dorsal ostioles. Notes. This is one of the more extreme pseudococcid types. Even though it lacks dorsal ostioles in any of the developmental stages, the presence of trilocular pores and tubular ducts of a type characteristic of the family make its assignment here seem entirely justified.

Paludicoccus distichlium (Kuwana) Kuwana Brown Mealybug (Figure 78) Pseudolecanium disticlium Kuwana, 1902. Ent. News 13:134-135, illus. Sphaerococcus disticlium (Kuwana), Fernald, 1903. Mass. Agr. Expt. Sta. Spec. Bull. 88, p.85.

212

TAXONOMY

OF

SPECIES

Fig. 78. Paludicoccus distichlium (Kuwana), collected on salt grass, Distichlis spicata (Gramineae), near Palo Alto (in salt marsh), Santa Clara County, California.

TAXONOMY Paludicoccus distichlium (Kuwana), Ferris, 1953. Atlas of Scale Insects of North America, Vol. VI: 392-393, illus. Type Locality and Host. Near Palo Alto (in salt marsh), Santa Clara County, California, on salt grass, Distichlis spicata (Gramineae). North American Distribution. California and Texas. Additional Hosts. Recorded only on species of Distichlis. External Features and Habitat. According to the original description, this mealybug is ellipsoidal in form, usually flat; pinkish-brown in color, shiny, the posterior segment darker color, and thinly coated with white, cottony wax. It occurs beneath the leaf sheaths of the stem and rootstock of grasses {Distichlis spp.), which grow only in salt marshes and saline areas. Recognition Characters. Adult female, mounted, 2.90 to 4.10 mm long, 1.00 to 1.50 mm wide; body form narrowly ovoid, posterior regions and lateral areas strongly sclerotized as far forward as head. Cerarii absent. Trilocular pores scattered rather sparsely over entire dorsum in midregion. Dorsal multilocular disk pores quite numerous, situated esspecially in lateral areas extending entirely about body, concentrated about spiracles. Tubular ducts about equal in diameter to central ring of a multilocular disk pore, extremely few, situated mainly on last abdominal segment, and along head margin. Body setae all very small, few in number. Anal ring situated slightly more than its own diameter from posterior apex of abdomen, ring quite strongly sclerotic, its pores few; its 6 setae scarcely half as long as diameter of ring. Venter with multilocular disk pores quite numerous, occurring especially along lateral areas extending entirely about body, especially concentrated about spiracles. Trilocular pores very sparsely distributed. Large and very diffuse cluster of minute circular pores present near lateral margin posterior to metathoracic spiracles. Tubular ducts of same type as those on dorsum occurring in small clusters especially in lateral areas of last 3 abdominal segments, also in marginal row along head. Body setae extremely sparse, all very small. Spiracular plates ovoid, quite large, strongly sclerotized. Circulus absent. Legs absent. Antennae usually 3segmented, rarely 4-segmented.

PHENACOCCUS Phenacoccus Cockerell, 1893. Ent. News. 4:317-318. Paroudablis Cockerell, 1900. Entomologist 33:85-87 (new synonymy). Phaenococcus Lindinger, 1942. Beiträge zur Fauna Perus Bd. 2, p. 115 (all but 6 copies destroyed in the war; reissued in 1952 as Bd. 3 [not 2], pp. 112-122.

OF

SPECIES

213

Notes. This is a very unusual kind of grass-infesting mealybug. It is included in this family only by the presence of trilocular pores and tubular ducts which are of the type that occurs in other forms of pseudococcids, and by the character of the antennae of the first-stage nymph. This monotypic species is without dorsal ostioles in any stage. The cerarii, circulus, and legs are also absent in the adult female. The spelling of the specific name, disticlium, is that designated by the original describer, and this has been corrected in accordance with the International Rules of Nomenclature. California Records (Map 74) San Diego Co.: San Felipe Creek, 1939, on Distichlis sp. (G. F. Ferris, UCD). Santa Clara Co.: Palo Alto, XII-18-01, on Distichlis spicata (I. Kuwana, UCD); VIII-17-17, on Distichlis spicata (G. F. Ferris, UCD).

COCKERELL Peukinococcus Sulc, 1944. Acta Soc. Sci. Nat. Morav. 16(11) :2-3, illus. Phenacoccus Cockerell, Ferris, 1950. Atlas of scale insects of North America, Ser. V; 120. Phenacoccus Cockerell, Ferris, 1953. Ibid., Vol. VI; 359-396. Phenacoccus Cockerell, Williams, 1962. Bull. Brit.

214

TAXONOMY

OF

SPECIES

Mus. (Nat. Hist.) Ent. 12 (1):32,34. Phenacoccus Cockerell, De Lotto, 1964. Ibid. 14 (8):365-366. Ferris (1950) remarks that "This genus is a very large group with, at present, an indeterminate number of species, which occur in North America, Europe, and Asia." Genotype. Pseudococcus aceris Signoret. Recognition Characters. Pseudococcidae with moderately oval body form, as mounted. Two pairs of ostioles. Normally with 18 pairs cerarii, there being a pair between frontal and ocular pairs, at times some may disappear from thoracic region. At times multilocular disk pores present. Tubular ducts usually present, rarely with an oral rim. Anal ring apical, of the usual "cellular" structure. Venter normally with multilocular disk pores, at times absent, never arranged in groups having a central duct. Quinquelocular pores usually present, although at times absent. Tubular ducts of the oralcollar type usually present. Circulus present or absent; if present, there at times being 2 or even 3. Legs well formed; claw with denticle or tooth on plantar surface. Antennae normally 9-segmented, although occasionally 8-segmented or less. Notes. This genus has for many years been used to include all species possessing 9-segmented antennae and a denticle or tooth on the claw. As understood, it included Puto (=Ceroputo Sulc) and certain other species, which are here placed in other genera. More recent developments indicate that the claw denticle, although still generally quite helpful in defining members of the genus Phenacoccus, cannot be completely relied upon as exemplifying this group alone. In the case of many species of Chorizococcus McKenzie and Spilococcus Ferris, this tiny denticle or tooth on the claw is present, and it occurs in combination with other characters not at all typical of the Phenacoccus series. As here limited, Phenacoccus seems—at least as far as the North American components are concerned—to form a quite natural group with no seriously disharmonious elements. Four new California species of Phenacoccus are described in this book. To the genus there are now assigned 35 species, 26 of which occur in California. During the course of the present investigations, and as a result of extensive survey and collecting of mealybugs throughout the state, it has become increasingly apparent that certain pseudococcid species show a range in variation of morphological characteristics far beyond that normally expected. This has been especially true for certain components of the Phenacoccus group and has resulted in the synomizing of 4 species previously considered to be quite distinctive. These species are: Phenacoccus advena McKenzie, P. harbisoni Peterson, P. milleri McKenzie, and P. nonarius McKenzie.

KEY Species of Phenacoccus in North America: Adult Females

1.

With more than 1 circulus 2 With 1 circulus or none 3 2(1). Abdominal cerarii with numerous conical setae, in clusters dearnessi King Abdominal cerarii for most part with but 2 conical setae each aceris Signoret 3(1). Ventral multilocular disk pores absent on last 3 or 4 abdominal segments . . 4 Ventral multilocular disk pores present, at least in region of vulva . . . . 7 4 ( 3 ) . Anal lobe cerarius with from 6 to 16 enlarged setae of uniform size, borne on a sclerotized area 5 Anal lobe cerarius with only 2 enlarged setae 6 5(4). Ventral multilocular disk pores present in midregion immediately posterior to circulus forward to prothorax megaulus McKenzie Ventral multilocular disk pores absent in area immediately posterior to circulus forward to prothorax eriogoni Ferris 6(4). Ventral quinquelocular pores present in midregion of thorax and on head; antennae 9-segmented destitutus McKenzie Ventral quinquelocular pores absent on thorax and head; antennae 7- or 8segmented lycii (Ferris) 7(3). Multilocular disk pores present on both dorsum and venter, at least on abdomen 8 Multilocular disk pores confined to venter 20 8(7). Circulus absent artemisiae Ehrhorn Circulus present 9 9 ( 8 ) . Ventral quinquelocular pores present on thorax 4 12 Ventral quinquelocular pores absent on thorax, except for perhaps 6 or 8 around mouthparts 10 10(9). Anal lobe cerarius with 4 to 7 moderately large conical setae of approximately the same size cajonensis McKenzie Anal lobe cerarius normally with 2, rarely with 3, conical setae of approximately the same size 11 11(10). Ventral oral-collar tubular ducts present 4 In Phenacoccus eremicus Ferris, the ventral quinquelocular pores are definitely present but are extremely small and inconspicuous, so much so that they are often difficult to find even in well-stained specimens.

TAXONOMY

12(9).

13(12).

14(13).

15(12).

16(15).

17(15).

18(17).

on thorax; occurs on various hosts alleni McKenzie Ventral oral-collar tubular ducts absent on thorax; to date, known only on "ocotillo," Fouquieria splendens (Fouquieriaceae) . . infernalis McKenzie Dorsum of thoracic region with considerable numbers of multilocular disk pores scattered over surface 13 Dorsum of thoracic region without multiocular disk pores, except that occasionally 1 or 2 pores may occur in this area 15 Ventral multilocular disk pores numerous in a broad band along lateral area of prothorax forward to eyes; occurring mainly on the maple family, Aceraceae acericola King Ventral multilocular disk pores absent in lateral area of prothorax, except that occasionally 1 to 4 pores may occur in this area 14 Anal lobe cerarius with from 6 to 10 enlarged conical setae of approximately same size, borne in a distinct sclerotized area; omnivorous plant feeder helianthi (Cockerell) Anal lobe cerarius with 2 paired conical setae, 3 or 4 smaller auxiliary setae, no basal area of sclerotization; occurring principally on the members of the grass family, Gramineae graminosus McKenzie At least one abdominal segment, except for ninth, with a median dorsal cerarius consisting of 2 or 3 enlarged setae and a few associated trilocular pores 16 Abdominal segments anterior to ninth without such a cerarius 17 Dorsal body setae throughout of the same shape and of approximately the same size as the cerarian setae; median dorsal cerarii on all abdominal segments except ninth flaveolus (Cockerell) Most of dorsal body setae distinctly shorter and more slender than the cerarian setae; median dorsal cerarii present only on abdominal segments 7 and 8 jranseriae Ferris With at least a few ventral multilocular disk pores on thorax, these usually situated in proximity of posterior spiracles and in submarginal area of metathorax hurdi McKenzie Multilocular disk pores absent on thorax venter 18 Multilocular disk pores on dorsum of abdomen from segment 8, forward to at least posterior margin of segment 5, and often present along sub-

19(18).

20(7). 21(20).

22(21).

23(22).

24(22).

25(24).

26(25).

OF

SPECIES

215

margin anterior to segment 5 19 Multilocular disk pores on dorsum of abdomen present mostly along posterior border of segment 8 forward to segment 6, absent on segment 5 and forward (except that occasionally 1 or 2 pores may occur in this area) eremicus Ferris Anal lobe cerarius with 7-9, usually 8, conical setae of approximately the same size multisetosus McKenzie Anal lobe cerarius with only 2 or 3, usually 3, conical setae of approximately the same size . . . . gossypii Townsend and Cockerell Circulus present 21 Circulus absent 32 Dorsum without tubular ducts except strictly marginal ones 22 Dorsum with a considerable number of tubular ducts scattered over surface 28 With 17 or 18 recognizable pairs of cerarii 24 With 15 recognizable pairs of cerarii or fewer 23 Dorsum with cerarii on last 4 or 5 abdominal segments, 2 - 4 on prothorax and head; antennae normally 9-segmented giganteus McKenzie Dorsum with cerarii present on all abdominal segments, 5-7 on thorax and head; antenae normally 8-segmented, infrequently 7-segmented echeveriae McKenzie With small but distinct sclerotized area surrounding base of each cerarius . . rubivorus Cockerell Cerarii without sclerotized area 25 Usually with only a few multilocular disk pores, these normally confined to posterior margin of eighth abdominal segment immediately anterior to vulva and ninth segment in area posterior to vulva defectus Ferris Usually with considerable numbers of multilocular disk pores around vulva and forward at least to seventh abdominal segment 26 Antennae normally 9-segmented; circulus moderately large, flaccid, and subject to distortion, sometimes slightly produced laterally; multilocular disk pores usually confined to area immediately around vulva and to segment 7; a large species, majority of specimens 4 - 5 mm. long solenopsis Tinsley Antennae normally 8-segmented; circulus normally quite small, circular or slightly oval, situated slightly above

216

TAXONOMY

27(26).

28(21).

29(28).

30(29).

31(30).

32(20).

33(32). 34(33).

OF

SPECIES

intersegmental line; multilocular disk pores on venter, from apex of abdomen forward to posterior margin of segment 5; species of average size, 3.00-3.50 mm long 27 Legs stout, robust with swollen femora and tibiae; hind tibiae with conspicuous translucent pores arranged in bands or clusters over most of segment, numbering from 118-135 (average 125.6); cerarian pairs variable with 14 to, at most, 17 pairs . . wilmattae Cockerell Legs moderately slender; hind tibiae comparatively slender to slightly swollen, translucent pores quite small and inconspicuous, scattered over segment, range from 44 to 71 (average 57.4); cerarii basically complete, normally 18 pairs, infrequently 17 solani Ferris Abdominal segments 7 and 8 each with a median dorsal cerarius with setae nearly as large as those of lateral cerarii; tubular ducts especially abundant in dorsolateral areas of abdominal segments 6 to 9 . . . . capensis Ferris Abdominal segments without median dorsal cerarii; tubular ducts of dorsum not thus concentrated 29 Many of dorsal body setae as stout and thick as setae of lateral cerarii, sometimes longer . . . . celtisifoliae Hollinger Dorsal body setae noticeably shorter and more slender than setae of lateral cerarii 30 With 17 or 18 recognizable pairs of cerarii colemani Ehrhorn With not more than 15 recognizable pairs of cerarii 31 Quinquelocular pores present on venter of head and thorax . . . minimus Tinsley Quinquelocular pores absent on venter of head and thorax . . dicoriae McKenzie With 18 recognizable pairs of cerarii on dorsum; venter with scarcely more than 10 multilocular disk pores, these immediately anterior to vulva pauperatus Ferris With 17 recognizable pairs of cerarii or fewer, these occurring on consecutive segments forward from apex of abdomen; multilocular disk pores on venter of ninth abdominal segment and forward at least to posterior margin of seventh 33 Octolocular pores scattered over both body surfaces . . . . incomptus McKenzie Octolocular pores absent 34 With at least a few oral-collar tubular ducts of same size scattered on dorsum of abdomen; antennae normally

9-segmented, rarely 8-segmented . . eschscholtziae McKenzie Dorsal oral-collar tubular ducts absent on abdomen, except for an enlarged submarginal one situated on each side of ninth segment usually anterior to anal lobe cerarius; antennae 6- to 8segmented lotearum McKenzie

Phenacoccus alleni McKenzie Allen Mealybug (Figures 79 and 80) Phenacoccus alleni McKenzie, 1964. Hilgardia 35( 10) :243-245, illus. Phenacoccus nonarius McKenzie, 1964. Ibid. 35(10) :253-254, illus. (new synonymy). Type Locality and Host. East Morongo Valley, San Bernardino County, California, on Hymenoclea salsola (Compositae). North American Distribution. Known only from California. Additional Hosts. Franseria dumosa (Compositae), Petalonyx thurberi (Loasaceae), and Sphaeralceaorcutti (Malvaceae). External Features and Habitat. This mealybug is heavily dusted with a gray, waxy secretion, so much so that segmentation is not visible. The body form is oval; body color light gray; body fluid exuded through both anterior and posterior ostioles also gray in color; anal ring secretion clear, capable of being ejected a distance of at least four inches; dorsum of abdomen with 2 very faint longitudinal lines; legs reddish. Small filaments around entire body margin, length undertermined. A filamentous ovisac is produced which is approximately one and one-half times as long as adult female. The first instar nymphs are orangish-pink and have no waxy secretion. The mealybug is found on the foliage of its host in the spring and summer and on the roots and crowns during the winter. The adult females seem to prefer the woody crowns, while the immatures prefer the roots. Ants were observed tending the mealybug. Recognition Characters. Adult females, mounted, 2.85 to 4.25 mm long, 1.60 to 2.25 mm wide; body form broadly oval. Dorsum normally with 18 pairs of cerarii, either without basal sclerotization or, at times, those on last 3 or 4 segments with small amount of sclerotization. Each cerarius normally with 2 conical setae, no auxiliary setae, slight concentration trilocular pores. Anal lobe cerarius and certain cerarii on head each may have 1-3 additional conical setae, slightly smaller than the 2 principal setae. Median dorsal cerarius on ninth abdominal segment present or absent; if present, usually consisting of 2 conical setae of same size and shape as those of lateral cerarii. Multilocular disk pores on dorsum variable, with extremes either restricted to abdominal segments 8-6 (see Figure 81), or present on all segments, except last, those on segments 3 and 4 confined to submar-

TAXONOMY

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Fig. 79. Phenacoccus alleni McKenzie, collected on Hymenoclea salsola (Compositae), at east Morongo Valley, San Bernardino County, California.

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Fig. 80. Phenacoccus alleni McKenzie, collected on Hymenoclea salsola (Compositae), at east Morongo Valley, San Bernardino County, California. Note especially fewer dorsal multilocular disk pores restricted to abdominal segments 8-6.

ginal areas, absent on thorax and head (see Figure 80). Trilocular pores evenly beset over entire dorsal surface, each pore appearing irregularly broken, with numerous fine lines. Minute circular pores scattered over dorsum. Considerable numbers of oral-collar tubular ducts scattered over entire dorsum, these infrequently with modified oral rims. Body setae short and slender. Anal ring apical, with no unusual characters; each of its 6 setae about twicé as long as greatest diameter of ring. Venter with considerable numbers of multilocular disk pores, variable, present on last 5-7 abdominal segments, normally extending across segments to body margin. Quinquelocular pores absent. Trilocular pores distributed fairly evenly. Minute circular pores scattered over venter. Oral-collar tubular ducts of 2 sizes in considerable numbers over entire venter, these infrequently with modified oral rims. Body setae slender and generally longer than those of dorsum. Circulus variable, sometimes compressed and probably divided by intersegmental line, or broadly oval and apparently not divided by this line. Legs comparatively large; hind femur with 0-12 translucent pores near distal end; hind tibia with translucent pores extremely variable (range from 8-51), scattered along most of segment; claw with prominent denticle. Antennae 9-segmented,5 moderately long. Notes. This species is closely related to Phenacoccus infernalis McKenzie, but differs primarily in having oral-collar tubular ducts and minute circular pores present on dorsum and venter of thorax; and in occurring on various hosts. On the other hand, P. infernalis lacks oral-collar tubular ducts and minute circular pores on dorsum and venter of thorax; and occurs only on "ocotillo," Fouquieria splendens (Fouquieriaceae). Prior to the description of P. alleni in 1964, a single specimen of this species, collected from the west side of Salton Sea, Imperial County, California, on Petalonyx sp. (Loasaceae), was placed by Ferris ( 1950) in his atlas as an extreme variant of P. eremicus Ferris. California Records (Map 75) Imperial Co.: Dixieland (near Seeley), III—1—65, on Franseria dumosa (R. A. Flock and L. Pineda, CDA); Westmorland, 11-23-65, on Sphaeralcea orcutti (R. Waegner and R. A. Flock, CDA) ; Salton Sea. Ill—1939, on Petalonyx sp. (G. F. Ferris, UCD). Inyo Co.: Townes Pass, VI-12-63, on Hymenoclea salsola (foliage) (D. R. Miller, UCD). Riverside Co.: 6 miles south Valerie Jean, VI-1063, on Petalonyx thurberi (D. R. Miller, UCD); 18 miles west Blythe, 1-29-65, on Franseria dumosa (D. R. Miller, UCD). San Bernardino Co.: 20 miles east Barstow, V 24-35, on Hymenoclea salsola (J. D. Maple, UCD); East Morongo Valley, IV-11-62, on Hymenoclea salsola (R. P. Allen, CDA); 24 miles south Twentynine Palms, IV-14-35, on Franseria dumosa (J. D. Maple, UCD). 5 Misinterpreted as 8-segmented antennae in original description.

Phenacoccus artemisiae Ehrhorn Artemisia Mealybug (Figure 81) Phenacoccus artemisiae Ehrhorn, 1900. Canad. Ent. 32:313, illus. Phenacoccus artemisiae Ehrhorn, Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., pp. 56-57, illus. Phenacoccus artemisiae Ehrhorn, Ferris, 1950. Atlas of Scale Insects of North America, Ser. V: 129130, illus. Type Locality and Host. Stevens Creek Canyon, near Mountain View, Santa Clara County, California, on California sagebrush, Artemisia californica (Compositae). North American Distribution. Known only from California. Additional Hosts. Artemisia spp. (Compositae) and Lotus sp. (Leguminosae). External Features and Habitat. The adult female slightly dusted with a fine, waxy secretion, is pruinose, pale sage-green, and lacks noticeable marginal tassels. The species is oviparous, the ovisac comparatively long and slender. This mealybug occurs on leaves, small twigs, crown, and roots of its host. It is often associated with ants. Recognition Characters. Adult female, mounted, 2.00 to 3.00 mm long, 1.00 to 2.40 mm wide; body form broadly oval. Dorsum normally with 18 pairs of cerarii—in early adult females borne on more or less distinct prominences—these eventually disappear with expansion of body, and remain only on head and posterior abdominal segments. Anal lobes prominent, noticeably pro-

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Fig. 81. Phenacoccus artemisiae Ehrhorn, collected on California sagebrush, Artemisia californica (Compositae), at Stevens Creek Canyon, near Mountain View, Santa Clara County, California.

TAXONOMY truding, somewhat pointed. Anal lobe cerarius with 2 quite slender setae, 1 or 2 small auxiliary setae, almost no concentration of trilocular pores. Remaining cerarii with 2, somewhat smaller setae, except in head region where there may be 3 setae, all with very few associated trilocular pores. Dorsum with multilocular disk pores arranged in a band along posterior margin of abdominal segments 8-6, those on segment 6 being concentrated toward lateral margin. Trilocular pores distributed fairly evenly. Minute circular pores scattered. Short, tubular ducts with a slight oral collar sparsely situated over dorsum. Body setae all smaller than cerarian setae, although similarly shaped. Anal ring apical, with no unusual characters; each of its 6 setae about twice as long as greatest diameter of ring. Venter with numerous multilocular disk pores in bands, mostly along posterior border of abdominal segments from vulva to segment 4, these more or less arranged in disconnected groups. Trilocular pores rather evenly beset on venter, except for certain "clear" areas on thorax. Minute circular pores scattered. Quinquelocular pores, smaller than multilocular pores, rather sparsely distributed in median region of anterior abdominal segments and on thorax and head. Tubular ducts somewhat smaller than those on dorsum, sparsely distributed over abdomen, very few on thorax. Body setae slender, mostly longer than those on dorsum. Circulus absent. Legs rather short; hind tibia with translucent pores scattered over most of segment; claw with distinct denticle or tooth on plantar surface. Antennae 8- or 9-segmented. Notes. The absence of a circulus and the presence of dorsal multilocular disk pores on abdomen readily distinguishes this species from other North American components of this group. This species appears to show considerable variation in the number of dorsal multilocular disk pores on abdomen, and in the number of recognizable cerarii—these often tending to be fewer than in the typical form. At the moment, there is little foundation for critical judgment in the matter of variability in structural details, especially quantitative, in mealybug species. California Records (Map 7 6 ) California (Sepse), VI-1910, on Artemisia californica (E. O. Essig, CAS). Inyo Co.: Antelope Springs, V I - 2 4 - 6 4 , on Artemisia sp. (crown and roots) (D. R. Miller, U C D ) ; Westgard Pass, V - l - 6 2 , on ? (A. A. Grigarick and J. L. Reed, U C D ) . Lassen Co.: 18 miles south Ravendale, V I - 1 4 - 6 2 , on Artemisia tridentata, associated with ants, Crematogaster sp. (R. R. Snelling, C D A ) ; 8 miles west Susanville, V I I - 8 - 6 4 , on Artemisia sp. (above ground under bark) (D. R. Miller, UCD). Los Angeles Co.: Whittier, 11-10-35 and X I I 1934, on Artemisia californica (J. D. Maple, U C D ) . Modoc Co.: 14 miles northwest Alturas (Big Sage Reservoir), V I - 2 7 - 6 3 , on Artemisia tridentata (on roots associated with ants Crematogaster coarctata coarctata) (D. R. Miller, U C D ) ; 7 miles east Cedar-

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ville, V I - 2 8 - 6 3 , on Artemisia tridentata (roots) (D. R. Miller, UCD) ; Cedarville road, V - 2 1 - 6 2 , on Artemisia tridentata (T. R. Haig, C D A ) ; 5.4 miles east Likely, V - 2 3 - 6 2 , on Artemisia tridentata (T. R. Haig, CDA). Nevada Co.: Russell Meadows, V I - 2 6 - 6 4 , on Artemisia sp. (roots) (D. R. Miller, UCD). Orange Co.: San Clemente, X I - 2 9 - 6 3 , on Artemisia californica (D. R. Miller, U C D ) ; 4 miles east El Toro, 111-25-64, on Artemisia sp. (crowns and roots) (D. R. Miller, UCD). Riverside Co.: Riverside, 1-16-35, on Artemisia californica (J. D. Maple, U C D ) ; 11-1935, reared in insectary (J. D. Maple, U C D ) . San Diego Co.: 5 miles east Escondido, X I I - 2 7 63, on Lotus sp. (crowns and roots) (D. R. Miller, U C D ) ; 1 mile south Pala, V I - 2 0 - 6 4 , on Artemisia sp. (D. R. Miller, U C D ) ; San Marcos, I I - 1 - 6 4 , on Artemisia californica (crowns and roots, associated with ants) (D. R. Miller, U C D ) . Santa Clara Co.: Stevens Creek, V I I I - 1 2 - 1 7 , on Artemisia californica (G. F. Ferris, UCD). Sierra Co.: 2 miles north Sagehen Creek, V I I - 6 64, on Artemisia sp. (on crown, under bark, and in Cerambycid borings) (D. R. Miller, UCD).

Phenacoccus cajonensis McKenzie, new species Cajon Mealybug (Figures 82 and 8 3 ) Type Locality and Host. Type and paratype adult females on flat-top, Eriogonum fasciculatum var. foliolosum (Polygonaceae), associated with ant, For-

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Fig. 82. Phenacoccus cajonensis McKenzie, new species, collected on flat-top, Eriogonum fasciculatum variety foliolosum (Polygonaceae), south of Cajon, San Bernardino County, California.

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Fig. 83. Phenacoccus cajonensis McKenzie, new species, collected on Eriogonum sp. (Polygonaceae), 7 miles east El Toro, Orange County, California. Note extreme variation of multilocular disk pores on both surfaces of body as compared to those in Figure 83.

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mica pilicornis, 2 miles south Cajon, San Bernardino County, California, collected June 11, 1963, by D. R. Miller. Additional paratype adult females collected in California on the crown and roots of various species of Eriogonum (Polygonaceae) are available as follows: on Eriogonum sp., Mountain Springs, Imperial County, collected June 21, 1964, by D. R. Miller; on Eriogonum sp., Whittier, Los Angeles County, collected December 30, 1934, and January 1, 1935, by J. D. Maple; on Eriogonum sp., 7 miles east El Toro, Orange County, collected June 11, 1964, by D. R. Miller and J. A. Froebe; on flat-top, Eriogonum fasciculatum, Irvine Lake, Orange County, collected November 29, 1963, by D. R. and J. L. Miller; on flat-top, Eriogonum jasciculatum, 5 miles north Modjeska Canyon, Orange County, collected November 29, 1963, by D. R. and J. L. Miller; on Eriogonum sp., 7 miles east San Juan Capistrano, Orange County, June 22, 1964, by D. R. Miller; on Eriogonum sp., associated with ants, at Silverado Canyon, Orange County, collected May 10, 1964, by D. R. Miller and J. A. Froebe; on Eriogonum sp., at Riverside, Riverside County, collected January 16, 1935 and February 22, 1935, by J. D. Maple; on Eriogonum sp., 7 miles east Mentone, San Bernardino County, collected July 30, 1964, by D. R. Miller and J. A. Froebe; on Eriogonum sp., at Red Hill, San Bernardino County, collected January 17, 1935, by J. D. Maple; on Eriogonum sp., 2 miles east Bonsall, San Diego County, collected June 20, 1964, by D. R. Miller; on Eriogonum sp., associated with ant, Crematogaster coarctata vermiculata, 5 miles southeast Fallbrook, San Diego County, collected May 9, 1964, by D. R. Miller; on Eriogonum sp., 1.4 miles east Boulevard, San Diego County, collected June 21, 1964, by D. R. Miller; and on Eriogonum sp., 1 mile northwest Valley Center, San Diego Co., collected June 23, 1964, by D. R. Miller. Type Material. Holotype adult female (single specimen on slide) and female paratypes in museum, University of California, Davis. Paratypes in United States National Museum, Washington, D. C., and in the collection of California State Department of Agriculture, Sacramento. North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. Adult female body covered with a smooth, gray, powdery, waxy secretion giving the body a gray appearance; with 1 or 2 pairs of minute, caudal wax filaments, sometimes absent. The legs are reddish; and the body contents when crushed are yellow. A filamentous, elongate ovisac is formed, covering all of adult female except the head. This mealybug was collected under rocks and on roots of its host, sometimes associated with ants. Recognition Characters. Adult female, mounted, 2.30 to 4.00 mm long, 1.40 to 2.40 mm wide; body form broadly oval. Dorsum normally with 18 pairs of cerarii. Anal

lobe cerarius with from 4 to 9 somewhat lanceolate conical setae, 3 or 4 of which are usually larger than others, few accompanying trilocular pores, borne on slightly sclerotized area. Penultimate cerarii normally with 2, rarely 3, setae, few trilocular pores, no sclerotization; remaining cerarii usually with 2 setae, except in anterior and head region where there may be 3-4, these somewhat smaller than those of anal lobes. Numerous dorsal multilocular disk pores in bands along posterior margin of all abdominal segments, except the last, rarely reduced to 1 or 2 on abdomen; extremely variable on thorax and head, ranging from 0-66 (average 16.9), when present generally most noticeable along body margin. Trilocular pores evenly beset over entire dorsal surface. Oral-collar tubular ducts fairly numerous on abdomen, scattered along submargin of thorax and head. Body setae fairly numerous, all short and slender. Anal ring apical, with no unusual characters; each of its 6 setae about twice as long as greatest diameter of ring. Venter with numerous multilocular disk pores on all abdominal segments forward to third segment, and usually in groups of variable numbers along lateral margin forward to head, single pore infrequently on head. Quinquelocular pores absent, except for perhaps 6 or 8 around mouthparts. Trilocular pores distributed fairly evenly except for certain "clear" areas in sternal region. Oral-collar ducts of same size and form as on dorsum present on abdomen, thorax and head. Body setae slender and generally longer than those of dorsum. Circulus present, moderately large, often produced laterally. Legs comparatively large; claw with prominent denticle or tooth on plantar surface. Antennae 9-segmented, moderately slender. Notes. This species appears to be quite specific on Eriogonum spp. (Polygonaceae). It is rather closely related to Phenacoccus helianthi (Cockerell), but differs principally in the absence of ventral quinquelocular pores, except for perhaps 6 or 8 around mouthparts, and in having an unsclerotized penultimate cerarius, with usually 2, rarely 3, setae. On the other hand, P. helianthi possesses ventral quinquelocular pores especially on anterior abdominal segments and in sternal region, penultimate cerarius is sclerotized, and usually with cluster of from 5 - 1 2 setae. California Records (Map 77) Imperial Co.: Mountain Springs, V I - 2 1 - 6 4 , on Eriogonum sp. (crown and roots) (D. R. Miller, UCD). Los Angeles Co.: Whittier, 1 - 1 - 3 5 and X I I 30-34, on Eriogonum sp. (J. D. Maple, U C D ) ; Red Hill, 1 - 1 7 - 3 5 , on Eriogonum sp. (J. D. Maple, UCD). Orange Co.: 7 miles east El Toro, V I - 1 1 - 6 4 , on Eriogonum sp. (D. R. Miller and J. A. Froebe, U C D ) ; Irvine Lake, X I - 2 9 - 6 3 , on Eriogonum jasciculatum (D. R. and J. L. Miller, U C D ) ; 5 miles north Modjeska Canyon, X I - 2 9 - 6 3 , on Eriogonum fasciculatum (D. R. and J. L. Miller, U C D ) ; Silverado Canyon, V - 1 0 - 6 4 , on Eriogonum sp. (crown

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and roots, associated with ant, Crematogaster coarctata vermiculata) (D. R. Miller and J. A. Froebe, UCD). San Bernardino Co.: 2 miles south Cajon, V I 11-63, on Eriogonum fasciculatum var. joliolosum (roots, associated with ants, Formica pilicornis) (D. R. Miller, U C D ) ; 7 miles east Mentone, VII-30-64, on Eriogonum sp. (crown and roots) (D. R. Miller and J. A. Froebe, U C D ) . San Diego Co.: 2 miles east Bonsall, VI-20-64, on Eriogonum sp. (D. R. Miller, U C D ) ; 1.4 miles east Boulevard, VI-21-64, on Eriogonum sp. (D. R. Miller, U C D ) ; 5 miles southeast Fallbrook, V - 9 - 6 4 , on Eriogonum sp. (crowns, associated with ant, Crematogaster coarctata vermiculata) (D. R. Miller, U C D ) ; 1 mile northwest Valley Center, VI-23-64, on Eriogonum sp. (crown and roots) (D. R. Miller, UCD).

Phenacoccus celtisifoliae Hollinger Celtis Mealybug (Figure 84; Color Plate III) Phenacoccus celtisifoliae Hollinger, 1923. Mo. Agr. Expt. Sta. Res. Bull. 58, pp. 47-48, illus. Phenacoccus picrospinus Ferris, 1950. Atlas of scale insects of North America, Ser. V: 156-157, illus. Phenacoccus celtisifoliae Hollinger, Ferris, 1953. Ibid., Vol. VI: 397. Type Locality and Host. Along Hinkson Creek south of Columbia, Boone County, Missouri, on hackberry, Celtis sp. (Ulmaceae).

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North American Distribution. California and Missouri. Additional Hosts. Lotus sp. (Leguminosae) and Photinia (= Heteromeles) arbutifolia (Rosaceae). External Features and Habitat. Adult female reported as light canary-yellow, covered with a heavy, white, mealy powder; legs, antennae, and mouth parts brownish or yellowish; body margined with very short, white masses of waxy secretion and a slight ridge of the same down the dorsum. When the insect is boiled in potassium hydroxide, it turns yellowish and then reddish brown. The mealybug infests the leaves and stems of its host (see col. pi. I I I ) . Recognition Characters. Adult female, mounted, 2.00 to 3.10 mm long, 1.10 to 2.00 mm wide; body form broadly oval. Dorsum normally with 18 pairs of cerarii. Anal lobe cerarius with 2 slender, sharply pointed, larger setae, 1 or more slender, but at times longer, auxiliary setae, several associated trilocular pores. Remaining cerarii each with but 2 setae, except ocular cerarius which may have 3-4, these are as long as those of anal lobe but somewhat more slender. Trilocular pores evenly beset over entire dorsal surface. Dorsum with small numbers of moderately large, oral-collar type ducts, these with rather heavy oral collar, situated in a single row across each abdominal segment, scattered over cephalic and thoracic areas, 1 or more just in from each of most cerarii. Body setae of various sizes, some extremely minute, others as large and long as those of lateral cerarii, if not even longer. Anal ring apical, with no unusual characters; each of its 6 setae about twice as long as greatest diameter of ring. Venter with rather small number multilocular pores in area about vulva forward to sixth segment. Extremely small and inconspicuous quinquelocular pores occupy median areas of thoracic segments, few appear on fifth abdominal segment and on segments anterior to this. Trilocular pores evenly beset on venter, except for certain "clear" areas on thorax. Considerable numbers of oral-collar tubular ducts with very slight oral collar situated in median and lateral areas of abdominal segments, few appearing in thoracic regions; all somewhat smaller than dorsal ducts. Body setae fairly numerous, generally longer than those on dorsum; lateral areas of abdomen each with several slender setae of various sizes. Circulus quite large, apparently oval, distorted in all available specimens. Legs noticeably small and moderately stout; hind tibia with translucent pores at distal end of segment; claw with prominent denticle. Antennae 9-segmented. Notes. The most obvious feature of this mealybug is the unusual length and slenderness of the cerarian setae and of some of those on dorsum, together with numerous long and slender setae in the ventral lateral areas of the abdomen. These characters, and a larger circulus, readily distinguish it from P. colemani. California Records (Map 78) Inyo Co.: Independence, Mt. Whitney Fish Hatch-

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Fig. 84. Phenacoccus celtisifoliae Hollinger, collected on Photinia (= Heteromeles) arbutifolia (Rosaceae), near Dunnigan, Yolo County, California. (Illustration prepared from specimens collected on Photinia [= Heteromeles] arbutifolia [Rosaceae], Orange County Park, Orange County, California.)

TAXONOMY ery, VIII-2-64, on Compositae (in Cerambycid tunnels) (D. R. Miller and J. A. Froebe, UCD). Orange Co.: Orange County Park, IV-14-36, on Photinia (= Heteromeles) arbutifolia (J. D. Maple, UCD). Ventura Co.: Fillmore, II-8-36, on Lotus sp. (J. D. Maple, UCD). Yolo Co. : Near Dunnigan, VII-28-64, on Photinia (= Heteromeles) arbutifolia (T. Kono and H. H. Keifer, CDA).

Phenacoccus colemani Ehrhorn Coleman Mealybug (Figures 85 and 86; Color Plate I)

Phenacoccus colemani Ehrhorn, 1906. Canad. Ent. 37:332-333. Phenacoccus colemani Ehrhorn, Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., pp. 57-58, illus. (misidentification). Phenacoccus colemani Ehrhorn, Ferris, 1950. Adas of scale insects of North America, Ser. V: 133134, illus. Phenacoccus tibiaegracilis McKenzie, 1960. Hilgardia 29(15):722-725, illus. (new synonymy). There have been numerous misidentifications under this name. Type Locality and Host. On Pascadero Road, south of Palo Alto, Santa Clara County, California, on Rubus sp. (Rosaceae). North American Distribution. California, Florida, Oregon, Puerto Rico, and Utah. Additional Hosts. Arctium lappa (Compositae), Arctostaphylos rudis (Ericaceae), Castilleia sp.

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(Scrophulariaceae), Encelia farinosa (Compositae), Eriogonum sp. (Polygonaceae), Garrya sp. (Garryaceae), Lantana camara (Verbenaceae), Onagraceae, Mahonia sp. (Berberidaceae), Pedicularis densiflora (Scrophulariaceae), Phacelia sp. (Hydrophyllaceae), Rubus vitifolius (Rosaceae), Symphoricarpos albus, and S. racemosus (Caprifoliaceae). It is quite commonly found beneath rocks associated with ants. The following species of ants have been found associated with this mealybug: Cremastogaster sp. (lineolatal), Formica subpolita camponoticeps, Formica subpolita Mayr (fusca group), and Formica spp. External Features and Habitat. Ehrhorn (1906) states the following about this species: "Adult female salmon pink, thinly covered with mealy secretion about 2XA mm long and IV2 mm wide. Legs and antennae light brown, eyes black. Female with egg sac about 5V2 mm long. Sac loosely woven of greenish-tinged cotton, not dense enough to hide the lemon-coloured eggs." Ferris (1950) states that this mealybug is "Thinly dusted with waxy powder, without noticeable lateral or caudal tassels. Oviparous, the eggs laid in a very loose, fluffy sac occurring on the leaves. It is apparently a form of spring and early summer, all the specimens at hand having been collected from late April to late May." This mealybug occurs on leaves, stems, and roots of its host. Ferris (1950) records it from beneath a rock in company with ants. He also reports a wingless male was taken in company with one of these collections. This has been substantiated by a more recent collection of the species under lava rock in close association with ants (see col. pi. I ) . These specimens were found under lava rock, closely associated with ants, near Hat Creek Post Office, Shasta County, California. In this case, when disturbed, the ants would immediately carry the mealybugs, waxy sacs and all, into ground cracks. Recognition Characters. Adult female, mounted, 1.70 to 2.50 mm long, 1.20 to 1.40 mm wide; body form broadly oval to strongly ovoid. Dorsum normally with 18 pairs of cerarii. Anal lobe cerarius with 2 rather slender, conical setae, 1 or 2 very small auxiliary setae, very slight concentration of trilocular pores. Remaining cerarii with 2 setae, except those on head each usually with 3 or 4 setae, these becoming extremely small along sides of body, accompanied by but 3 or 4 pores. Trilocular pores sparsely beset on dorsal surface. Minute circular pores scattered. Oral-collar type ducts occur over body as far forward as head, arranged in a transverse row of scarcely more than 10 across most abdominal segments, also in small groups of scarcely more than 4-5 or singly near each of the cerarii. These ducts have quite distinct oral collar, although they vary in size, being very small and inconspicuous in some specimens. Dorsal body setae few, all very small. Anal ring apical, with no unusual characters; each of its 6 setae about twice as long as greatest diameter of ring. Venter with relatively few multilocular disk pores

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Fig. 85. Phenacoccus colemani Ehrhorn, collected on Rubus sp. (Rosaceae), on Paseadero Road, south of Palo Alto, Santa Clara County, California.

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Fig. 86. Phenacoccus colemani Ehrhorn, collected on Garry a sp. (Garryaceae), Cuyama, Ventura County, California. Note especially the long, slender legs; relatively large, flaccid circulus; and rather numerous ventral quinquelocular pores. These characteristics seem to fall well within the limits of variation for the species.

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in region of vulva, occasionally a very small number as far forward as fifth abdominal segment. Quinquelocular pores variable, either somewhat restricted to head region near mouthparts or scattered from sixth abdominal segment forward. Trilocular pores distributed fairly evenly except in thoracic region where there may be certain "clear" areas. Minute circular pores scattered. Tubular ducts present in small numbers across median region of abdominal segments and lateral areas of these segments, these all without oral collar, smaller than dorsal ducts. A few oral-collar tubular ducts same size and type as those on dorsum occur in lateral areas of abdomen and thoracic region. Body setae, slender and comparatively sparse, some noticeably longer than those on dorsum. Circulus variable in shape, either small, oval, undivided, with heavily sclerotized perimeter, or quite large, flaccid, apparently divided, perimeter unsclerotized. Legs well developed, variable; tibiae normally with translucent pores scattered almost full length; claw with very small denticle on plantar surface. Antennae 9-segmented, rather short. Notes. This species is perhaps most closely related to Phenacoccus solani Ferris, but differs from it mainly in the possession of oral-collar tubular ducts on the dorsum, these structures absent in the species last mentioned. In some specimens of P. colemani, these ducts are very small, but they are definitely present in all examples here considered to belong to this species. California Records (Map 79) Alameda Co.: Oakland, 111-20-22, in "ants nest" (E. C. Van Dyke, UCD); Redwood Peak, 111-19-22, in "ants nest" (E. C. Van Dyke, CAS). Lassen Co.: Doyle, VII-22-65, on Eriogonum sp. (T. R. Haig, CDA). Marin Co.: Cypress Ridge, IV-2-22, in "ants nest" (J. Lamiman, CAS). Mendocino Co.: Mendocino, VIII-6-58, in "soil associated with ants" (J. R. Heifer, UCD). Modoc Co.: Cedarville Road, V-21-62, "under rock"—tended by ants, Formica subpolita camponoticeps (T. R. Haig, CDA); 3 miles south east Lava Beds National Monument, VI-29-63, on Onagraceae and Phacelia sp. (roots) (D. R. Miller, UCD). Nevada Co.: 1 mile south Truckee, X-16-63, "under rock" with ants, Formica sp. (T. R. Haig, CDA). Placer Co.: 13 miles east Foresthill, VI-1962, "under rock" with ants, Lasius flavus (R. R. Snelling, CDA). Plumas Co.: 2 miles east Chilcoot, 111-31-64, "under rock" associated with ants (T. R. Haig, CDA); Graeagle, VI-18-64, "under rock" (T. R. Haig, CDA). Riverside Co.: Palm Springs, III-9-44, on Encelia farinosa (T. D. A. Cockerell, UCD). San Mateo Co.: Bear Gulch, IV-28-17, on Pedicularis densifiora (G. F. Ferris, UCD); North of Woodside, IV-7-17, "under rock" in nest of Crematogaster sp. (lineolatal) (G. F. Ferris, UCD); Woodside, V-10-17, on Rubus vitifolius (G. F. Ferris,

UCD); VII-19-17, on Castilleia sp. (G. F. Ferris, UCD). Santa Barbara Co.: Lompoc (Vandenberg Village), VI-26-63, on Arctostaphylos rudis (crown) (H. L. McKenzie, UCD). Santa Clara Co.: South of Palo Alto (on Pescadero Road), VI-5-1900, on Rubus sp. (collector ?, UCD); Stanford University (San Francisquito Creek). 11-15-17, (G. F. Ferris, UCD); V-7-17, on Symphoricarpos racemosus (G. F. Ferris, UCD); I V 14-17, "under rock" with ant, Formica sp. (G. F. Ferris, UCD). Shasta Co.: 10.7 miles north Hat Creek Post Office, V-23-63, "under lava rock" in association with ants, Formica subpolita Mayr (fusca group) (T. R. Haig, CDA); 6 miles south Hat Creek Post Office, V 23-63, "under lava rock" (T. R. Haig, CDA); Jet. Highways 44 and 89, 1 mile south Hat Creek, IV22-64, "under rock" with ant, Formica sp. (T. R. Haig, CDA). Sierra Co.: 5 miles north Floriston, III—31—64, "under rock" associated with ants (T. R. Haig, CDA). Siskiyou Co.: Lava Beds National Monument, Merill Cave, VI-29-63, on Onagraceae (roots associated with ants, Formica subpolita camponoticeps) (D. R. Miller, UCD). Ventura Co.: Near Cuyama, VI-11-59 and I X - 3 59, on Garrya sp. (M. Cravens and J. Schall, CDA).

Map 79. Distribution in California of Phenacoccus colemani Ehrhorn.

Sa/i Francit co SanMatto

TAXONOMY Phenacoccus

dearnessi King

Two-Circuli Mealybug (Figure 87) Phenacoccus dearnessi King, 1901. Canad. Ent. 33:180. Phenacoccus cockerelli King, 1903. Ibid. 35:195. Phenacoccus betheli Cockerell, 1912. Ibid. 44:301. Phenacoccus regnillhoi Hollinger, 1923. Mo. Agr. Expt. Sta. Res. Bull. 58, pp. 49-51. Phenacoccus dearnessi King, Ferris, 1950. Atlas of scale insects of North America, Ser. V: 135-136, illus. Phenacoccus dearnessi King, Ferris, 1953. Ibid., Vol. VI: 397. Type Locality and Host. London, Ontario, Canada, on thorn, Crataegus sp. North American Distribution. Arizona, California, Canada, Colorado, Michigan, Missouri, and Texas. Additional Hosts. Amelanchier pallida, Amelanchier sp. (Rosaceae), Prunus (= Amygdalus) havardii (Rosaceae). External Features and Habitat. This mealybug is a very rotund species. Its dorsum has transverse areas of thin secretion which do not conceal the dark red color, and the margins are equipped with a very short fringe of tassels which more or less flow together. The species apparently comes to maturity in midsummer, judging by available collection dates. The species occurs primarily on the twigs and stems of its host. Recognition Characters. Adult female, mounted, 1.75 to 3.50 mm long, 1.25 to 3.10 mm wide; body shape almost circular. Dorsum with 9-10 pairs of cerarii counting forward from anal lobes. Anal lobes cerarius with 5-10 short, stout, slightly lanceolate setae, these all about same size, no auxiliary setae, accompanied with few trilocular pores set in slightly sclerotized area. Remaining abdominal cerarii contain even greater number of similar setae, these forming a patch on side of body. Anterior to abdomen these patches become irregular, toward head scarcely distinguished as distinct cerarii. Dorsum with numerous trilocular pores; no multilocular disk pores; apparently no tubular ducts. Scattered sparsely over dorsum are body setae similar in form, and in part equal in size to those of cerarii. Anal ring apical, with no unusual features; its 6 setae about twice as long as greatest diameter of ring. Venter with very few multilocular disk pores, these concentrated in region of vulva forward to posterior margin of sixth abdominal segment. Small quinquelocular pores scattered in midregion of anterior abdominal segments forward to head. Trilocular pores less numerous than on dorsum, noticeable "clear" areas showing in sternal area. A few small, oral-collar tubular ducts situated especially on posterior segments of abdomen and in lateral areas. Two circuli present, these circular or slightly oval, quite small, anterior one being smaller than posterior. Legs well developed, not especially stout; claw with distinct denticle or tooth on plantar surface. Antennae normally 9-segmented, quite short.

OF

SPECIES

231

Notes. The large patches of conical setae forming the cerarii, and the 2 small circuli easily differentiate this species from all other components of this group. The accompanying illustrations of P. dearnessi were made by Ferris from specimens collected on Prunus ( = Amygdalus) harvardii from the Chisos Mountains, Texas. A reexamination of these specimens show slightly larger legs and perhaps fewer ventral multilocular disk pores than indicated in Ferris' illustrations. California Records (Map 80) Alpine Co.: Markleeville, VIII-11-22, on Amelanchier sp. (E. Bethel, UCD); Pleasant Valley, VII28-61, on Amelanchier pallida (R. F. Wilkey, CDA).

Imperfect Mealybug

(Figure 88) Phenacoccus defectus Ferris, 1950. Atlas of scale insects of North America, Ser. V: 137-138, illus. Type Locality and Host. Permanente Creek, Santa Clara County, California, on Eriophyllum confertiflorum (Compositae). North American Distribution. Found only in California. Additional Hosts. Ambrosia sp. (Compositae), Baccharis ? sp. (Compositae), Chenopodiaceae, Echeveria longissima (Crassulaceae), Echeveria spp. (Crassulaceae), Eriogonum umbellatum var. polyanthum (Polygonaceae), Eriophyllum confertiflorum (Compositae), Euphorbia sp. (Euphorbiaceae), Gramineae, Helianthus sp. (Compositae), Lupinus sp. (Leguminosae), Mortardella sp. (Labiatae), Phacelia sp. (Hydrophyllaceae), Sempervivum tectorum (Crassulaceae), "succulent," Tetradymia spinosa (Compositae), and "under rock."

232

TAXONOMY

OF

SPECIES

Fig. 87. Phenacoccus dearnessi King, collected on Amelanchier pallida (Rosaceae), at Pleasant Valley, Alpine County, California. (Illustration prepared from specimens collected on Prunus [= Amygdalus] harvardii [Rosaceae], Chisos Mountains, Texas.)

TAXONOMY

Fig. 88. Phenacoccus defectus Ferris, collected on Eriophyllum positae), at Permanente Creek, Santa Clara County, California.

OF

SPECIES

confertiflorum

(Com-

233

234

TAXONOMY

OF

SPECIES

External Features and Habitat. No available information on external features of this species. The mealybug occurs on the foliage, crown, and roots of its host. Recognition Characters. Adult female, mounted, 2.20 to 3.50 mm long, 1.40 to 2.50 mm wide; body form broadly oval. Dorsum normally with 18 pairs of cerarii. Anal lobe cerarius with 2 rather slender, slightly lanceolate setae accompanied by a single rather long, slender seta or variable numbers of small auxiliary setae, few scattered trilocular pores. Other cerarii with, for most part, 2 smaller and more slender lanceolate setae and 3 or 4 trilocular pores. Dorsum very sparsely beset with trilocular pores. Tubular ducts absent. Dorsal body setae few and scattered, all slightly lanceolate, for most part smaller than smallest cerarian seta, an occasional seta as large as those of lateral cerarii; seventh and eighth abdominal segments with median pair of at times slightly enlarged setae, these placed rather close together, scarcely meriting designation of dorsal cerarii. Anal ring apical, with no unusual features; each of its 6 setae about twice as long as greatest diameter of ring. Venter with very few multilocular disk pores situated around vulva, number of pores not exceeding total of 20 in specimens examined. Quinquelocular pores apparently absent. Trilocular pores rather evenly distributed. Very few—perhaps scarcely more than 20—small, inconspicuous oral-collar tubular ducts scattered over posterior portion of abdomen. Bod}' setae, for the most part, slightly longer than those on dorsum. Circulus present, oval, normally extending across intersegmental line between segments 4 and 5, rarely not divided by this line. Legs well developed; claw with slight denticle or tooth on plantar surface. Antennae normally 9-segmented, infrequently 8-segmented. Notes. According to Ferris (1950), "This species was erroneously included in Phenacoccus colemani Ehrhorn by Ferris in 1918, but is definitely not that species." It is practically a "twin" of P. pauperatus Ferris, but differs in possessing a circulus. It is also closely related to P. solani Ferris, except that the ventral multilocular disk pores are very few and usually confined to last two abdominal segments; it has a 9-segmented antennae; and it normally has a larger circulus which is divided by the intersegmental line between abdominal segments 4 and 5. P. solani, on the other hand, usually has more ventral multilocular disk pores, these distributed on last 3-5 abdominal segments; possesses an 8-segmented antenna; and normally has a small circular or oval circulus not transversely divided by intersegmental line across which it lies. California Records (Map 81) California (quarantined at Honolulu, Hawaii), I I 13-50, on Sempervivum tectorum (H. Makino, USNM). Alameda Co.: Berkeley (U.C. Botanical Gardens),

XII-1948, on Echeveria longissima (R. E. Beer, USNM); Berkeley, 1-24-49, on Echeveria rubella and E. scheeri (W. S. Sibray, CDA ) ; VI-26-62, on Echeveria sp. (P. Manichote, UCD). Alpine Co.: Winnemuca Lake, VII-14-64, on undetermined plants (roots) and Gramineae (roots) (J. A. Froebe, UCD). Butte Co.: Gridley, IV-30-59, on Ambrosia sp. A. G. Forbes and R. F. Wilkey, CDA). Inyo Co.: Bishop, VI-13-63, on Tetradymia spinosa (foliage) (D. R. Miller, UCD). Modoc Co.: 5.4 miles east Likely, V-23-62, "under rock" (R. F. Wilkey and T. R. Haig, CDA). Riverside Co.: 2 miles south Coachella, VI-10-63, on Chenopodiaceae (foliage) (D. R. Miller, UCD); Indio, XI-8-62, on "succulent" (N. W. Getz and Bascom, CDA); 1 mile north White Water, 1-25-65, on Euphorbia sp. (on crown just below ground) (D. R. Miller, UCD). San Bernardino Co.: 1 mile west Joshua Tree, X I 30-63, on Phacelia sp. (crowns) (D. R. Miller, UCD); Morongo Valley, VI-9-63, on Baccharis ? sp. (foliage) (D. R. Miller, UCD); 15 miles south Twentynine Palms, IV-14-35, on Lupinus sp. (J. D. Maple, UCD). San Diego Co.: Ramona, 1-16-57 and II—1-57, on Echeveria sp. (G. L. Hill, CDA). Santa Clara Co.: Permanente Creek, V-5-17, on Eriophyllum confertiflorum (G. F. Ferris, UCD); Stevens Creek, VIII-26-17, on Monardella sp. (G. F. Ferris, UCD). Siskiyou Co.: Valentine Cave, Lava Beds National Monument, VI-26-63, on Eriogonum umbellatum var. polyanthum (roots and crown) (D. R. Miller, UCD). Ventura Co.: Fillmore, 11-27-36, on Helianthus sp. (J. D. Maple, UCD).

Map 81. Distribution in California of Phenacoccus

defectus Ferris.

Mann-*SmI

FrafKitco San Mat*«

TAXONOMY

Phenacoccus destitutus McKenzie, new species Destitute Mealybug (Figure 89) Type Locality and Host. 5 miles south Cedarville, Modoc County, California, on Eriogonum latifolium var. saxícola (Polygonaceae), June 28, 1963, by D. R. Miller. Type Material. Holotype adult female (single speciment on slide) in collection of museum, University of California, Davis. . .North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. A single adult female of this species was found intermixed with large numbers of specimens of Phenacoccus eriogoni Ferris. Unfortunately, no field observations were made of its external features since it was so completely camouflaged by the other mealybugs. The mealybug was collected on the roots of its host. Recognition Characters. Holotype adult female, mounted, 2.00 mm long, 1.20 mm wide; body form oval. Dorsum normally with 18 pairs of marginal cerarii. Anal lobe cerarius with 2 enlarged conical setae, 2 or 3 auxiliary setae, 1 of which is usually larger than others, few accompanying trilocular pores, no sclerotized area. Remaining cerarii usually with 2 setae, slight group trilocular pores. Submarginal band of modified cerarii with 1 or 2 setae much smaller than those of marginal cerarii, situated from eighth abdominal segment forward to anterior thorax; similar type modified cerarii in median area from eighth abdominal segment forward to second. Trilocular pores rather evenly beset on dorsum, each pore internally shattered. Minute circular pores scattered over surface. Tubular ducts absent. Body setae short and slender. Anal ring apical, with no unusual characters; each of its 6 setae about twice as long as greatest diameter of ring. Venter destitute of multilocular disk pores. Quinquelocular pores present in considerable numbers from immediately above vulva forward in midregion of abdomen, thorax, and head. Trilocular pores similar to those on dorsum, fairly evenly beset on venter except for certain "clear" areas in sternal region. Minute circular pores scattered on surface. Oral-collar tubular ducts of 2 sizes, few in number, the largest ones most abundant, situated in submarginal row from eighth abdominal segment to metathorax; 3-4 smaller ducts on abdominal segments 8-7, absent elsewhere on venter. Body setae slender, some noticeably longer than those on dorsum. Circulus present, distinctly oval, situated on fourth abdominal segment, not divided by segmental line. Legs comparatively large; tibiae with light scattering of small translucent pores along entire segment; claw with denticle or tooth on plantar surface near tip. Antennae 9-segmented, slender.

OF

SPECIES

235

Notes. This species is somewhat related to Phenacoccus lycii (Ferris) but differs mainly in having numerous ventral quinquelocular pores in midregion of thorax and on head, and 9-segmented antennae. P. lycii, on the other hand, has ventral quinquelocular pores restricted to abdomen, and 7- to 8-segmented antennae. California Records Map (82) Modoc Co.: 5 miles south Cedarville, VI-28-63, on Eriogonum latifolium var. saxicola (roots) (D. R. Miller, U C D ) .

Phenacoccus dicoriae McKenzie Dicoria Mealybug (Figure 90) Phenacoccus dicoriae McKenzie, 1961. Hilgardia 31(2):28-30, illus. Type Locality and Host. 3 miles west Garnet, Riverside County, California, on Dicoria canescens (Compositae). North American Distribution. Known only from California. Additional Hosts. Franseria dumosa (Compositae). External Features and Habitat. No available information at this time on external appearance of this species. The mealybug apparently occurs above ground on leaves and stems of its host. Recognition Characters. Adult female, mounted, 2.80 to 4.80 mm long, 2.00 to 3.00 mm wide; body form broadly oval. Dorsum with 11 to 15 recognizable pairs of cerarii, each normally with 2 cerarian setae, these generally

236

TAXONOMY

OF

SPECIES

Fig. 89. Phenacoccus destitutus McKenzie, new species, collected on Eriogonum latifolium variety saxicola (Polygonaceae), south of Cedarville, Modoc County, California.

TAXONOMY

OF

SPECIES

Fig. 90. Phenacoccus dicoriae McKenzie, collected on Dicoria canescens (Compositae), west of Garnet, Riverside County, California.

237

238

TAXONOMY

OF

SPECIES

quite small, often rather widely separated, especially those anterior to last 3 or 4 pairs, all without accompanying auxiliary setae, very slight concentration of trilocular pores. Slight suggestion of cerarii along midregion of body, in some specimens so inconspicuous that they are difficult to find. Trilocular pores abundant, evenly disposed over entire dorsum. Minute circular pores scattered. Oralcollar tubular ducts quite numerous, especially in lateral areas of abdomen, arranged segmentally on most abdominal segments, also situated submarginally on meso- and metathorax. Anal ring of normal form for genus; its 6 setae about twice as long as greatest diameter of ring. Venter with numerous multilocular disk pores, situated especially in midregion of abdomen from apical to fourth segments, absent on thorax and head. Quinquelocular pores entirely absent on body. Trilocular pores fairly numerous, evenly distributed on venter. Minute circular pores scattered. Oral-collar tubular ducts, with very slight oral collar, occur in considerable numbers in median and lateral areas of abdomen, few such ducts in thoracic region. Body setae slender, for most part, considerably longer than those on dorsum. Circulus moderately large, transversely oval. Legs proportionately well developed; claw with prominent denticle or tooth on plantar surface. Mouthparts relatively slender. Antennae 9-segmented, moderately long. Notes. This mealybug appears to resemble, at least to some degree, Phenacoccus minimus Tinsley, but differs from it in absence of ventral quinquelocular pores on thorax and head, these structures definitely present in the species last mentioned. Other differences include a greater number of ventral multilocular disk pores in P. dicoriae, as compared to fewer ventral multilocular disk pores in P. minimus. California Records (Map 83)

Map 83. Distribution in California of Phenacoccus dicoriae McKenzie.

Riverside Co.: 3 miles west Garnet, II-5-60, on Dicoria canescens (S. W. Brown and U. Nur, UCD). San Diego Co. : Borrego Spring, 1-27-65, on Franseria dumosa, (crown) (D. R. Miller, UCD).

Phenacoccus echeveriae McKenzie Echeveria Mealybug (Figure 91) Phenacoccus echeveriae McKenzie, 1960. Hilgardia 29(15) :715-717, illus. Phenacoccus echeveriae McKenzie, 1962. Ibid. 32(14):651. Type Locality and Host. Near Hemet, San Jacinto River, Riverside County, California, on perennial succulent, Echeveria sp. (Crassulaceae). North American Distribution. California and Nevada. Additional Hosts. Compositae, Lupinus sp. (Leguminosae). External Features and Habitat. No habitat information was included about this species collected on Echeveria. A subsequent collection of the mealybug, however, was made on the roots of Lupinus sp. No remarks were made in either collection relative to its external appearance. Recognition Characters. Adult female, mounted, 1.75 to 2.20 mm long, 1.00 to 1.30 mm wide; body form rotund. Dorsum with variable number of cerarii, in type species with at least 15 discernible pairs, or associations of "setae" with triloculars on each side that can be regarded as more or less developed cerarii. Anal lobe cerarius with 2 conical setae, 1 or more auxiliary setae, cluster trilocular pores. Penultimate cerarii without auxiliary setae, cerarian pair smaller, more slender than those on anal lobes. Cerarii on thorax with setae so slender and widely separated as to leave some question as to their validity. Dorsum with trilocular pores rather evenly distributed. Minute circular pores numerous, scattered over entire dorsal surface. A few oral-collar tubular ducts occur along margin of posterior abdominal segments. Dorsal body setae all small and slender. Anal ring with no unusual features; its 6 setae about twice as long as greatest diameter of ring. Venter with considerable numbers of multilocular disk pores, situated predominantly in midregion of abdomen, from apical to fourth abdominal segments. Trilocular pores generally distributed over entire venter. Minute circular pores numerous, scattered over entire ventral surface. Small oral-collar tubular ducts occur intermingled with multiloculars, in considerable numbers in lateral areas of ninth to second abdominal segments, as well as a group between anterior and posterior spiracles and body margin. Body setae slender, generally longer than those on dorsum. Circulus transversely oval, extending across fold between fourth and fifth abdominal segments. Legs relatively stout; hind coxae with cluster of translucent

TAXONOMY

OF

SPECIES

Fig. 91. Phenacoccus echeveriae McKenzie, collected on a perennial succulent, Echeveria sp. (Crassulaceae), near Hemet, San Jacinto River, Riverside County, California.

239

240

TAXONOMY

OF

SPECIES

pores at base; irregular cluster translucent pores on apical half hind tibiae; claw with small denticle or tooth near tip. Antennae normally 8-segmented, at times with 7-segments. Notes. This species resembles, at least to some degree, Phenacoccus minimus Tinsley, but differs in the absence of ventral quinquelocular pores on thorax and head, these structures present in the species last mentioned. Also there are only a very few small, dorsal, submarginal, oral-collar ducts present in P. echeveriae, whereas these structures in P. minimus are quite numerous over entire dorsum and are noticeably enlarged. Specimens collected 3 miles west of Morongo Valley in San Bernardino County on Compositae possess 8- or 9-segmented antennae and generally fewer ventral multilocular disk pores on abdomen, as compared to specimens in the type series. In all other respects, however, they agree in detail with what is currently recognized as P. echeveriae. California Records (Map 84) Riverside Co.: Near Hemet, San Jacinto River, 11-22-36, on Echeveria sp. (F. R. Piatt, C D A ) . San Bernardino Co.: 3 miles west Morongo Valley, VI-9-63, on Compositae (roots, associated with ants) (D. R. Miller, U C D ) .

Lauen

Map 84. Distribution in California of Phenacoccus echeveriae McKenzie.

San foneticoSan Matte

Phenacoccus eremicus Ferris Eremicus Mealybug (Figure 92) Phenacoccus eremicus Ferris, 1950. Atlas of scale insects of North America, Ser. V: 139-140, illus. Type Locality and Host. Near Mojave, San Ber-

nardino County, California, on burro-weed, Franseriadumosa (Compositae). North American Distribution. Known only from California. Additional Hosts. Encelia jarinosa, Encelia sp. (Compositae), Hymenoclea salsola (Compositae), Marrubium vulgare (Labiatae), Pluchea sericea (Compositae), and Sida hederacea (Malvaceae). The single adult female, on Petalonyx sp. (Loasaceae), west side of Salton Sea, Imperial County, California, mentioned by Ferris (1950) as possibly an extreme variant of P. eremicus is instead Phenacoccus alleni McKenzie. External Features and Habitat. The adult females of this mealybug are covered with a fine, powdery, white, waxy secretion. There is no evidence of anal or lateral filaments. The mealybug occurs on the crown and roots of its host. Recognition Characters. Adult female, mounted, 2.50 to 5.50 mm long, 1.00 to 4.00 mm wide; body form broadly oval. Dorsum normally with 18 pairs of cerarii. Anal lobe cerarius with 2 quite small, slightly lanceolate cerarian setae, 1 or 2 auxiliary setae of same type, accompanied by very slight concentration trilocular pores. Remaining cerarii with but 2 setae, except for 1 or 2 in head region which may have 4 or 5 setae, these all very small, inconspicuous, with very slight concentration of trilocular pores. In some specimens cerarii slightly indicated along midregion of body, although usually so inconspicuous that they are difficult to find. Dorsum with a somewhat variable number and arrangement of multilocular disk pores, these occurring for most part along posterior border of abdominal segments 6-7, in few cases, completely absent. Trilocular pores very numerous throughout. Minute circular pores scattered. Oral-collar tubular ducts, slightly larger in diameter than trilocular pores and with very slight oral collar, scattered on abdomen and along margin of thorax. Body setae very few, for most part very small, some equalling smaller cerarian setae in size, all somewhat lanceolate, scattered over body surface. Anal ring apical, with no unusual features; each of its 6 setae about twice as long as diameter of ring. Venter with multilocular disk pores extremely numerous in region of vulva, these extending forward in diminishing numbers to posterior margin of fourth abdominal segment, few situated in lateral areas of segments 6-7. Quinquelocular pores extremely small and inconspicuous, difficult to find even in wellstained preparations, situated on sixth abdominal segment forward to prothorax. Trilocular pores distributed fairly evenly. Minute circular pores scattered. Oral-collar ducts considerably numerous, these of same size and form as those on dorsum, becoming fewer or almost disappearing on head and thorax. Body setae slender and generally longer than those of dorsum. Circulus noticeably large, usually slightly oval, at times appearing somewhat produced laterally, this

TAXONOMY

OF

Fig. 92. Phenacoccus eremicus Ferris, collected on burro-weed, Franseria (Compositae), near Mojave, San Bernardino County, California.

SPECIES

dumosa

241

242

TAXONOMY

OF

SPECIES

probably due to distortion. Legs well developed; hind tibia with translucent pores scattered along most of segment; claw with distinct denticle or tooth on plantar surface. Antennae normally 9-segmented. Notes. This species is rather closely related to Phenacoccus alleni McKenzie, differing from it principally in possessing ventral quinquelocular pores on sixth abdominal segment forward to, but not including, head, and in lacking dorsal oral-collar tubular ducts on thorax and head. On the other hand, P. alleni lacks ventral quinqueloculars but possesses dorsal oral-collar tubular ducts on thorax and head. Phenacoccus eremicus is also closely related to Phenacoccus infernalis McKenzie, differing from it principally in having dorsal multilocular disk pores occurring for the most part along posterior margin of abdominal segments 6 and 7, whereas in P. infernalis these structures are present on dorsum of all abdominal segments, except the last. In some specimens of P. eremicus, the dorsal multiloculars seem restricted to segment 7 and to be very few or, in a few instances, completely absent. Specimens without evidence of dorsal multiloculars will key to Phenacoccus minimus Tinsley, although the dorsal oral-collar tubular ducts in P. eremicus are small, being slightly larger in diameter than the trilocular pores, and less abundant on thorax, whereas in P. minimus, these structures are relatively very large and more numerous in this area. Also, in P. eremicus the ventral quinqueloculars are extremely small and inconspicuous even in well-stained specimens, whereas in P. minimus these structures are well developed and conspicuous. California Records (Map 85)

Trinity j

Piuma,

Map 85. Distribution in California of Phenacoccus

> BolttS

eremicus Ferris.

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•

RinrùJ*

Imperial Co. : South of Brawley, 11-25-65, on Sida hederacea (R. A. Flock and G. T. Okumura, UCD); Calexico, 11-28-57, on Pluchea sericea (G. G. Beevor, CDA) ; 1 mile east Glamis, 1-24-64, on Encelia farinosa, Franseria dumosa, and Hymenoclea salsola (W. H. Lange and H. W. Michalk, UCD); 5 miles north Truckhaven, XII-31-63, on Encelia sp. (crown and roots) (D. R. Miller and J. A. Froebe, UCD). Kern Co.: Near Mojave, IV-26-36, on Franseria dumosa (G. F. Ferris, UCD). Los Angeles Co.: Near Lancaster (Antelope Valley), IV-27-36, on Franseria dumosa (G. F. Ferris, UCD). Riverside Co.: 3 miles west Garnet, II-5-60, on Compositae (U. Nur, UCD); Palm Springs (on road to Palm Canyon), 11-27-63, on Franseria dumosa (crown and roots) (H. L. McKenzie, UCD). San Bernardino Co.: 5.5 miles south Garlic Springs, IV-7-35, on Franseria dumosa (J. D. Maple, UCD); 4 miles south Kramer Junction, VI-24-64, on Franseria dumosa (in Cerambycid tunnels) (D. R. Miller, UCD); 24 miles south Twentynine Palms, IV-14-35, on Franseria dumosa (J. D. Maple, UCD). Ventura Co.: Fillmore, V-2-35, on Marrubium vulgare (J. D. Maple, UCD).

Phenacoccus eriogoni Ferris Cerarii Mealybug (Figure 93) Phenacoccus eriogoni Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., p. 58, illus. Phenacoccus eriogoni Ferris, 1950. Atlas of scale insects of North America, Ser. V: 141-142, illus. Type Locality and Host. Stevens Creek, Santa Clara County, California, on tibinagua, Eriogonum nudum (Polygonaceae). North American Distribution. Known only from California. Additional Hosts. Baccharis sp. (Compositae), Eriogonum deflexum brachypodium, E. latifolium var. saxicola, Eriogonum spp. (Polygonaceae), Lotus scoparius (Leguminosae), Peucephyllum sp. (Compositae), Phacelia ramosissima (Hydrophyllaceae), Pluchea sericea (Compositae), and Purshia glandulosa (Rosaceae). External Features and Habitat. This mealybug is reported as being thickly covered with waxy secretions, the lateral filaments short and broad, the caudal ones longer. An ovisac was not observed. Mature specimens have been taken in the field from May to October. The species is found on the foliage, crown, and roots of its host. Recognition Characters. Adult female, mounted, 2.00 to 4.00 mm long, 1.10 to 3.10 mm wide; body form broadly oval to quite rotund. Dorsum normally with 18 pairs of cerarii. Anal lobe cerarius with as many as 10 enlarged, slightly lanceolate, cerarian setae, these accompanied by but a few trilocular pores set in a faintly sclerotized area.

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Fig. 93. Phenacoccus eriogoni Ferris, collected on tibinagua, Eriogonum nudum (Polygonaceae), at Stevens Creek, Santa Clara County, California.

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Remaining cerarii with 3-8 somewhat smaller setae of same form as those on anal lobes, slight group trilocular pores. Trilocular pores on dorsum unusually large, almost equalling diameter of sockets about base of anal lobe setae. Minute circular pores scattered. Tubular ducts absent on dorsal surface. Body setae few, all much smaller than those of cerarii, except for small group present, on midline of eighth abdominal segment (rarely on seventh segment), this group including 3-4 setae of same size and form as those of anal lobes, accompanied by small cluster trilocular pores, presenting appearance of a genuine dorsal cerarius. Anal ring situated at posterior apex of body, with no unusual features; its 6 setae about twice as long as greatest diameter of ring. Venter completely destitute of modified multilocular. (quinquelocular?) disk pores in all available specimens, except, at times, 1-5 such pores near mouth parts. Trilocular pores about same size as those on dorsum, scattered over entire venter. Minute circular pores scattered. Very few oral-collar tubular ducts situated in region of vulva. Body setae small except for few long and slender ones in midregion of abdomen and sternum and on head. Circulus present, distinctly oval. Legs relatively very large; hind tibia with translucent pores along entire segment; claw with very prominent denticle or tooth on plantar surface. Antennae normally 9-segmented. Notes. This species is very closely related to Phenacoccus megaulus McKenzie, but differs mainly in the absence of dorsal oral-collar tubular ducts, as well as ventral modified multilocular (quinquelocular?) disk pores in sternal region, except for possibly an occasional such pore in this general area. P. megaulus, on the other hand, always possesses a variable number of relatively large oral-collar tubular ducts on dorsum, as well as numerous ventral modified multilocular (quinquelocular?) disk pores situated in median area of fifth abdominal segment forward to and including prothorax. California Records (Map 86) Butte Co.: Chico, VII-8-62, on Eriogonum sp. (T. R. Haig, CDA); Paradise, VI-5-62, on Eriogonum sp. (T. R. Haig, CDA). Inyo Co.: 5 miles east Big Pine, VII-18-63, on Eriogonum deflexum ssp. brachypodium (roots, crown, and foliage) (D. R. Miller, UCD). Nevada Co.: 6 miles northwest Nevada City, V-6-66, on Eriogonum sp. (R. F. Wilkey, CDA). Riverside Co.: 2 miles south Coachella, VI-10-63, on Pluchea sericea (foliage) (D. R. Miller, UCD); Indio, 111-26-38, on Baccharis sp. (roots) (R. C. Dickson, UCD); Riverside, VII-4-35, on Lotus scoparius (J. D. Maple, UCD); V-13-62, on Phacelia ramosissima (J. A. Powell, UCD). San Bernardino Co.: East Highlands, V-9-45, on Eriogonum sp. (R. C. Dickson, UCD). Santa Clara Co.: Stephens Creek, X-14-16, V-617, and VIII-27-17, on Eriogonum nudum (G. F. Ferris, UCD); V-5-17 and V-13-17, on Eriogonum sp. (G. F. Ferris, UCD).

Shasta Co.: Redding, X-16-62, on Eriogonum sp. (T. R. Haig, CDA). Sierra Co. : 5 miles east Floriston, VIII-26-65, on Eriogonum sp. (T. R. Haig, CDA). Siskiyou Co.: 8 miles north Yreka, IX-2-63, on Eriogonum sp. (J. S. Buckett, UCD). Tehama Co.: 7.5 miles west Dales, VI-25-63, on Eriogonum latifolium var. saxícola (roots, crown, and foliage) (D. R. Miller, UCD).

Phenacoccus eschscholtziae McKenzie California Poppy Mealybug (Figures 94 and 95; Color Plate II) Phenacoccus eschscholtziae McKenzie, 1961. Hilgardia 31(2):30-31, illus. Phenacoccus advena McKenzie, 1964. Ibid. 35(10): 241-243, illus. (new synonymy). Phenacoccus milleri McKenzie, 1964. Ibid. 35(10): 251-253, illus. (new synonymy). Type Locality and Host. Box Springs Canyon, near Shavers Well, Riverside County, California, on Eschscholtzia sp. (Papaveraceae). North American Distribution. California and Mexico. Additional Hosts. Agoseris ? (Compositae), Amsinckia sp. (Boraginaceae), Coreopsis californica (Compositae), Cryptantha angustifolia (Boraginaceae), Echeveria sp. (Crassulaceae), Eriogonum sp. (Polygonaceae), Erodium cicutarium (Geraniaceae), Helianthemum scoparium andersoni (Cistaceae), Layia glandulosa (Compositae), Malacothrix glabrata (Compositae), Nama demissum (Hydrophyllaceae), Plantago sp. (Plantaginaceae), Rumex hymeno-

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Fig. 94. Phenacoccus eschscholtziae McKenzie, collected on Eschscholtzia sp. (Papaveraceae), at Box Springs Canyon, near Shavers Well, Riverside County, California.

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Fig. 95. Phenacoccus eschscholtziae McKenzie, collected on chia, Salvia columbariae (Labiatae), west of Joshua Tree, San Bernardino County, California. Note especially the more numerous cerarii; ventral multilocular disk pores restricted to last three abdominal segments; and the much swollen hind tibia, as compared to the illustration of this species shown in Figure 94.

TAXONOMY sepalus (Polygonaceae), and Salvia columbariae (Labiatae). External Features and Habitat. In life this mealybug is of a somewhat lavender-gray color and is very thinly dusted with a waxy secretion. Apparently there are no lateral tassels, and the caudal tassels are very small or lacking. No ovisac was observed (see col. pi. II).

This species feeds on the crowns or underground portion of its host plant. Recognition Characters. Adult females, mounted, 2.10 to 3.90 mm long, 1.00 to 2.75 mm wide; body form broadly oval. Dorsum with cerarian pairs extremely variable, ranging from 3 to 15: 3 to 8 pairs on abdomen, 0 to 5 pairs on thorax, 0 to 2 pairs on head. Cerarii reduced and often difficult to discern because the conical setae are situated unusually far apart. Anal lobe cerarius with 2 large conical setae of variable shapes, often with 3 to 4 smaller conical setae, no auxiliary setae, very slight concentration trilocular pores, small amount basal sclerotization; remaining cerarii with 2 enlarged conical setae similar to anal lobe pair except progressively smaller anteriorly except for those in ocular pair, usually widely separated, often with no indication of closely associated trilocular pores. Trilocular pores evenly beset over entire dorsum. Minute circular pores scattered. Oral-collar tubular ducts, some with a very slight oral-rim, on abdomen in median area and along submargin, a few scattered on thorax and head. Body setae very small, sparse. Anal ring apical, with no unusual characters; each of its 6 setae nearly twice as long as greatest diameter of ring. Venter with multilocular disk pores variable in distribution, restricted to last 3, 4, or 5 abdominal segments, absent elsewhere, except that 1 specimen had a single pore on venter of fourth abdominal segment. Trilocular pores evenly beset. Minute circular pores scattered. Oral-collar ducts, of the same type as those on dorsum, found mostly on abdomen, scattered on thorax and head. Body setae slender, more numerous and mostly longer than those on dorsum. Circulus absent. Legs of moderate size; hind tibia variable in shape, with from 12 to 98 (average 47.1)° very small translucent pores scattered over most of segment; claw with well-developed denticle. Antennae moderately long, normally 9-segmented, occasionally 8 segmented, rarely 7-segmented, distal segment usually divided and of a characteristic shape—widest at the dividing line and tapering rather abruptly in both directions. Notes. This is an extremely variable mealybug, and for this reason two illustrations are here included to more adequately depict the species. Two species, Phenacoccus advena and P. milleri, both described by the writer in 1964, have been placed as synonyms of P. eschscholtziae McKenzie because of this variability. This much variation in morphological structures is quite unusual in the Pseudococcidae. 8 Based on a critical examination of 65 specimens from numerous hosts and localities.

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Phenacoccus eschscholtziae is perhaps most closely allied to Phenacoccus incomptus McKenzie, a Mexican species, but is easily distinguished from it by the absence of octolocular pores, whereas in the species last mentioned these pores are scattered over dorsal and ventral body surfaces. California Records (Map 87) I m p e r i a l C o . : Seeley, 1 1 1 - 2 5 - 6 5 , Plantago sp. (erectal) (R. A. Flock, C D A ) Inyo Co.: 5 miles east Big Pine, V I I - 1 8 - 6 3 , on Eriogonum deflexum ssp. brachypodium (roots, crowns, and foliage) (D. R. Miller, U C D ) ; 3 miles northwest Shoshone, X I I - 2 9 - 6 3 , on Cryptantha sp. (roots) (D. R. Miller, U C D ) . Orange Co.: Irvine Lake, V I - 6 - 6 3 , on Malacothrix saxatilis var. tenuifolia (roots, associated with ants, Crematogaster coarctata vermiculata) (D. R. Miller, UCD). Riverside Co.: Box Spring Canyon, near Shavers Well, I V - 1 4 - 3 5 , on Eschscholtzia sp. (J. D. Maple, U C D ) ; 8 miles west Palm Desert, 111-10-65, on Cryptantha angustifolia (H. L. and V. B. McKenzie, UCD). San Bernardino Co. : 1 mile west Joshua Tree, I V 9 - 6 3 and X I - 3 0 - 6 3 , on Agoseris(?) sp., Amsinckia sp., Coreopsis californica, Erodium cicutarium, Layia glandulosa, Malacothrix glabrata, Nama demissum, and Salvia columbariae (D. R. Miller, U C D ) ; 5 miles south Kramer Junction, 1 - 2 4 - 6 5 , on Amsinckia sp. (roots) (D. R. Miller, U C D ) , Twentynine Palms, X I I - 2 3 - 6 4 , on Eriogonum sp. (roots) (D. R. and J. F. Miller, U C D ) ; 2 miles west Yucca Valley, I V 9 - 6 3 , on Rumex hymenosepalus and Salvia columbariae (D. R. Miller, U C D ) . San Diego Co.: 0.5 miles north Oceanside, 1 - 2 9 64,"under rock" (D. R. Miller, U C D ) ; 2 miles northwest Valley Center, on Helianthemum scoparium andersoni (roots) (D. R. Miller, U C D ) . San Francisco Co.: San Fancisco, 11-20-13, in ants nest in sand hill (J. C. Bridwell, U C D ) .

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Phenacoccus giganteus McKenzie Giant Mealybug (Figure 96; Color Plate II) Phenacoccus giganteus McKenzie, 1964. Hilgardia 35(10):245-246, illus. Type Locality and Host. 12 miles south Shoshone, Inyo County, California, on burro-weed, Franseria dumosa (Compositae). North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. In life, the pinkish body shows faintly through the very thin dusting of a whitish secretion. There are no noticeable caudal or lateral tassels. Before oviposition, the adult female becomes enclosed in a rather tough sac ( formed of fine, waxy threads. Fully mature specimens were taken in April (see col. pi. II). This mealybug occurs on the crown or underground parts of its host. Recognition Characters. Adult females, mounted, 5.00 to 8.00 mm long, 3.25 to 5.50 mm wide; body form broadly oval. Dorsum with cerarii on the last 4 or 5 abdominal segments, occasionally 1 or 2 pairs distiguishable on prothorax and 1 or 2 ocular pairs on head. Anal lobe cerarius with a small area of sclerotization. Each cerarius with 2 conical setae and without auxiliary setae; in addition to the 2 large conical setae, each of the anal lobe cerarii and possibly the ocular cerarii may have 1 or 2 slightly smaller conical setae. All cerarii with a very slight concentration of triloculor pores. Trilocular pores numerous and evenly beset over entire dorsum. Minute circular pores scattered. No tubular ducts of any kind. Body setae short and slender. Anal ring apical, with no unusual characters; each of its 6 setae slightly longer than greatest diameter of ring. Venter with multilocular disk pores on all abdominal segments except second and third, especially numerous on the last 4 segments, lacking elsewhere. Trilocular pores evenly beset. Minute circular pores scattered. Oral-collar ducts fairly numerous on abdomen, especially along submargin, scattered elsewhere on venter. Body setae slender, slightly longer than those of dorsum. Circulus comparatively large, usually shaped like a loaf of bread, situated on fourth abdominal segment, not capable of folding along the intersegmental line. Legs well formed, comparatively small; hind tibiae with 62 to 172 translucent pores (average 98.6) distributed along entire segment; claw with prominent denticle. Mouthparts comparatively stout. Antennae 9-segmented, proportionately very small. Notes. One of the outstanding features of this species is its enormous size, which is the obvious reason for naming it giganteus. Even the smaller of the adult females are usually 5.00 mm long—the size of the largest individuals in many other known species. This mealybug appears related to Phenacoccus echeveriae McKenzie but has only 6 to 8 pairs of

cerarii and a 9-segmented antenna, as compared to the 15 pairs of cerarii normal for P. echeveriae and its 7- or 8-segmented antenna. In size, giganteus approaches Phenacoccus eremicus Ferris, but the absence of dorsal multiloculars on abdomen, and presence of minute circular pores on both body surfaces precludes confusion with that species. On the other hand, P. eremicus normally possesses dorsal multiloculars on sixth and seventh abdominal segments, and the minute circular pores are absent on both body surfaces. California Records (Map 88) Inyo Co.: 12 miles south Shoshone, IV-13-63, on Franseria dumosa (D. R. Miller, UCD). Riverside Co.: Desert Hot Springs, IV-15-65, on Franseria dumosa (crown and roots) (D. R. and J. F. Miller, UCD).

Phenacoccus gossypii Townsend and Cockerell Mexican Mealybug (Figures 97 and 98; Color Plate I) Phenacoccus gossypii Townsend and Cockerell, 1898. Jour. N.Y. Ent. Soc. 6:170-171. Phenacoccus helianthi var. gossypii Townsend and Cockerell, Cockerell, 1899. 111. Nat. Hist. Survey Bull. V:391. Phenacoccus colemani Ehrhorn, Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., pp. 57-58 (misidentification). Phenacoccus gossypii Townsend and Cockerell, Myers, 1928. Calif. Dept. Agr. Monthly Bull. 17 (6) :355-358, illus.

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Fig. 96. Phenacoccus giganteus McKenzie, collected on burro-weed, Franseria dumosa (Compositae), south of Shoshone, Inyo County, California.

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Fig. 97. Phenacoccus gossypii Townsend and Cockerell, collected on "beans" (Leguminosae), at Willows, Glenn County, California. (Illustration prepared from specimens collected on Phaseolus sp. [Leguminosae], at Palo Alto, Santa Clara County, California.)

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Fig. 98. Phenacoccus gossypii Townsend and Cockerell, collected on sunflower, Helianthus tiiveus (= H. techrodes) (Compositae), at Sand Hills, east of Gray's Well, Imperial County, California. Note circulus which is only slightly produced laterally. All other features agree in detail with our present concept of this species.

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Phenacoccus gossypii Townsend and Cockerell, Myers, 1932. Jour. Econ. Ent. 2 5 ( 4 ) : 8 9 1 - 8 9 6 , illus. Phenacoccus gossypii Townsend and Cockerell, Zimmerman, 1948. Insects of Hawaii, Univ. Hawaii Press, 5:161-165, illus. Phenacoccus gossyppi Townsend and Cockerell, Ferris, 1950. Atlas of scale insects of North America, Ser. V: 145-147, illus. Phenacoccus harbisoni Peterson, 1965. Pan-Pacific Ent. 4 1 ( 2 ) :96-100, illus. (new synonymy). Type Locality and Host. Presumably at Frontera, State of Tabasco, Mexico, host not specifically indicated in original description. It is assumed that this is included in the first reference to its occurrence on "a cultivated ornamental plant called 'amistad' which is closedly allied to cotton and is probably a species of Gossypium." North American Distribution. Arizona, California, Colorado, Cuba, Florida, Hawaii, Illinois, Louisiana, Maryland, Mississippi, New York, Puerto Rico, and Texas. Additional Hosts. Ferris (1950) stated: "The host list is so extensive that no attempt will here be made to recite it in full, since the species apparently may occur on almost any flowering plant." A few of these hosts are listed as follows: Althaea sp. (Malvaceae), Ambrosia sp. (Compositae), Calendula sp. (Compositae), Capsicum jrutescens longum (Solanaceae), Chrysanthemum sp. (Compositae), Citrus sp. (Rutaceae), Coleus sp. (Labiatae), Crotalaria sp. (Leguminosae), Dombeya sp. (Sterculiaceae), Erigeron sp. (Compositae), Erythrina spp. (Leguminosae), Geranium sp. (Geraniaceae), Gossypium sp. (Malvaceae), Hedera helix (Araliaceae), Heliconia sp. (Musaceae), Lantana sp. (Verbenaceae), Lycopersicon sp. (Solanaceae), Phaseolus limensis (Leguminosae), and Solanum spp. (Solanaceae). External Features and Habitat. In life this mealybug is of a grayish color and is covered with a thin waxy secretion. The lateral filaments are very short, and the caudal ones do not exceed one-fourth the body length. A firm, long, white ovisac is produced, this sinuous and often twice as long as the adult females (see col. pi. I ) . Although the species normally occurs above ground on the leaves and stems of the host, it may, at times, also be found feeding on the crown and roots. Recognition Characters. Adult female, mounted, 2.50 to 3.75 mm long; 1.40 to 2.10 mm wide; body form broadly oval. Dorsum normally with 18 pairs of cerarii. Anal lobe cerarius normally with 2-3 quite large, slightly lanceolate cerarian setae, often accompanied by 1 or more somewhat smaller auxiliary setae, slight concentration trilocular pores. Remaining cerarii usually with 2 setae (occasional head cerarii with 3 setae), slightly smaller than those of anal lobes, accompanied by small group trilocular pores. Multilocular disk pores present on dorsum in a band or row across posterior border of abdominal segments 8-4, few situated laterally on posterior part

of segment 3. Trilocular pores evenly beset over entire dorsum. Minute circular pores scattered on abdomen. Small numbers of oral-collar tubular ducts situated over entire dorsal surface. Body setae rather numerous, all about same shape as those of cerarii, variable in size, none larger than setae of lateral cerarii. Anal ring apical, with no unusual features; each of its 6 setae about twice as long as greatest diameter of ring. Venter with numerous multilocular disk pores, these situated in region posterior to vulva forward to third abdominal segment, also in lateral areas of segments 7-3. Numerous quinquelocular pores present in midregion of head, thoracic segments, anterior abdominal segments. Trilocular pores distributed fairly evenly except for certain "clear" areas in midregion of thorax. Very small oral-collar ducts, some with slight development of an oral rim, especially in deeply stained specimens, present in moderate numbers in median region of abdomen; other oral-collar ducts, same size as those on dorsum, occur in lateral abdominal regions, also in median and lateral regions of thoracic segments. Body setae slender, slightly longer than those of dorsum. Circulus relatively quite large, usually produced laterally into a narrow extension on each side, resembling an ox yoke, or with anterior margin only slightly produced laterally, not divided by intersegmental line. Legs well developed; hind tibia with translucent pores scattered along segment. Claw with prominent denticle or tooth on plantar surface. Antennae normally 9-segntented, slender. Notes. This is the so-called "Mexican mealybug" of economic entomologists. It is a very common pest of ornamentals of various kinds and has been reported as a minor pest of lima beans. The most characteristic morphological feature of this mealybug is the form of the circulus, this usually being relatively very large and produced laterally into a narrow extension on each side. Phenacoccus gossypii is related to P. multisetosus McKenzie, herein described as new, but differs in having 3 conical setae, 1 or 2 smaller auxiliary setae in anal lobe cerarius; and the circulus, while usually strongly produced laterally, is without a sclerotized process or spur arising on each side. P. multisetosus, on the other hand, has numerous, usually 8, conical setae, without auxiliary setae in anal lobe cerarius; and the circulus is comparatively small, transversely oval with a sclerotized process or spur arising on each side. In the character of possessing both dorsal and ventral multilocular disk pores on abdomen, it appears somewhat related to Phenacoccus graminosus McKenzie. It differs from the species last cited, however, in the characteristic shape of the circulus, already mentioned, and in the absence of dorsal multilocular disk pores on the thorax, head, and ventrally along submargin of thorax. In P. graminosus the circulus is quite small, laterally oval, and multilocular disk pores are present in considerable numbers on

TAXONOMY thorax and head, as well as ventrally along submargin of thorax. California Records (Map 89) California (quarantined at Honolulu, Hawaii), IV24-50, on Geranium sp. (D. F. Chong, USNM); VI-15-50, on Geranium sp. (H. Makino, USNM). Alameda Co.: Berkeley (U. C. Greenhouse), X 22-39, on Apium graveolens (Kolber, CAS); X - 4 39, on Conium maculatum (E. O. Essig, CAS); X 2-39, on Geranium sp. (leaves and stems) (C. Michener, CAS); Berkeley (quarantined in Yuba Co.), Ill—16—33, on Geranium sp. (H. A. Crane, CDA); IX—4-39, on Jacobinia pauciflora (E. O. Essig, CAS); X-20-39, on Veronica sp. (E. O. Essig, CAS); Oakland, X-8-28, on Taraxacum vulgare (B. Veblem, CAS); San Leandro, V-22-31, on Lantana sp. (L. S. Jones, CDA). Butte Co.: Chico, XI-2-38, on ? (W. B. Carter, CDA); Oroville, X-19-37, on Acanthus mollis (Fitzgerald, CDA); 1-11-30, on Choisya sp. (C. H. Kinsley, CDA). Contra Costa Co.: El Cerrito, VI-5-47, on Eschscholtzia sp. (G. F. Ferris, UCD); Pinole, X-3-31, on "lionstail" (J. B. Steinweden, CDA). Fresno Co.: Fresno, 1-12-39, on Euonymus sp. (J. R. Clark, CDA); VI-14-39, on Hibiscus sp. and Gardenia sp. (T. B. Gallion, CDA); 1-13-39, on Ixia sp. (J. R. Clark, CDA). Glenn Co.: Willows, VIII-31-31, on "beans" (C. H. Wren, CDA). Humboldt Co.: (Alderpoint quarantined from San Jose) X-2-33, on Geranium sp. (W. E. Peacock, CDA); Areata, X-31-34, on "miscellaneous plants" (W. E. Peacock, CDA); (Eureka quarantined from San Francisco) X-12-34, on "ornamental" (W. E. Peacock, CDA). Imperial Co.: Brawley, VII-13-65, on Distichlis sp. (R. A. Flock, CDA); Sand Hills near Gray's Well, VI-22-63, on Helianthus niveus (= H. tephrodes) (associated with ant, Myrmecocystus semirufus) (R. L. Westcott, UCD); near Gray's Well, XII-28-63, on Helianthus niveus (= H. tephrodes) and Petalonyx thurberi (D. M. Peterson, UCD). Kern Co.: Bakersfield, XII-8-28, on Verbena sp. (B. L. Fox, CDA). Kings Co.: Lemoore, V-30-27, on Tagetes sp. (L. E. Myers, LEM). Los Angeles Co.: Alhambra, IX-24-19, on Coprosoma sp. (A. D. Borden, UCD); IX-24-19, on Tagetes sp. (A. D. Borden, UCD); Claremont, VI-1924, on Citrus sp. (H. M. Armitage, CDA); Compton, V-25-26, on "ornamentals" (L. E. Myers, LEM); Covina, X-9-11, on Solanum douglasii (E. W. Rust, USNM); Durante, XII-21-25, on Citrus sp. "orange" (L. E. Myers, LEM); Glendale, 1923, on Calluna sp. (G. R. Gorton, UCD); 1-1920, on Geranium sp. (G. F. Ferris, UCD); La Verne, X I 28-26, on Chrysanthemum sp. (L. E. Myers, USNM); Long Beach, X-16-41, on Xanthium sp. (F. R. Piatt, LEM); Los Angeles, X-28-32, on Chrysanthemum sp. (T. F. Catching, USNM); XI1921, on Chrysanthemum sp. (H. J. Ryan, CDA);

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XI-1921, on Jacaranda sp. (H. J. Ryan, CDA); X I I 18-56, on Lantana sp. (N. L. H. Krauss, USNM); Monrovia, 1923, on Franseria sp. (E. Bethel, UCD); Montebello, XII-28-42, on Solanum melongena (M. Miller, LEM); Pasadena, 1-1921, on Abutilón sp. (R. S. Woglum, UCD); III-1920, on Abutilón sp. (R. S. Woglum, UCD); XII-1921, on Cupressus sp. (A. G. Smith, UCD); 111-25-27, on Hederá helix (L. E. Myers, LEM); IX-22-33, on Lantana sp. (J. B. Steinweden, CDA); VI-1-20, on Wyethia sp. (A. D. Borden, UCD); Pasadena (Huntington Estate), 1921, on Wisteria sp. (H. M. Armitage, CDA); Redondo Beach, VI-13-64, on Oenothera cheiranthifolia (D. M. Peterson, UCD); San Fernando, 1918, on Calinyction sp. (H. J. Quayle, UCD); Santa Monica, 1923, on Sonchus sp. and Malacothrix sp. (E. Bethel, UCD); Torrance, IX-1-20, on Lycopersicon esculent um var. commune (J. M. Wright, USNM); Van Nuys, X-1923, on "ornamentals" (collector ?, CDA); Whittier, 1922, on Solanum tuberosum (H. M. Armitage, UCD); 11-10-35, on Salvia sp. (J. D. Maple, UCD); 11-24-35, on ? (J. D. Maple, UCD); 111-10-35, on Lotus sp. (J. D. Maple, UCD). Marin Co.: San Rafael, XI-1-61, on XJlex europaeus (D. Maddox, UCD). Merced Co.: Arena ?, XI-19-40, on "fern" (M. J. Williams, CDA); Merced, VI-3-32, on Acacia sp. (D. P. Wheeler, CDA). Monterey Co.: Gonzales, X-15-60, on Geranium sp. (D. W. Berry, UCD); 5 miles north Marina, I I 8-64, on Eriophyllum confertiflorum (crowns) (D. R. Miller and J. A. Froebe, UCD) ; Monterey, IV30-53, on Lotus scoparius (R. P. Allen, CDA). Orange Co.: Costa Mesa, 11-24-32, on "lionstail" (L. E. Myers, LEM); Huntington Beach, 1-31-64, on Haplopappus sp. (roots and foliage) (D. R. Miller and J. A. Froebe, UCD); San demente, IX-8-42, on "miscellaneous plants" (G. G. Beevor, CDA); VIII-15-65, on Lotus scoparius (G. M. Harper, CDA); 9 miles east San Juan Capistrano, VI-22-64, on Diplacus sp. (crown and roots) (D. R. Miller, UCD); Santa Ana, VI-1920, on Acacia ? sp. (H. M. Armitage, CDA); 11-17-32, on Begonia sp. (T. E. Leod, LEM); III—29—26, on Geranium sp. (L. E. Myers, CDA); 1929, on Hibiscus syriacus (collector ?, CDA); West Santa Ana, IX-15-65, on "ornamental orange" (J. Clod and D. Park, CDA); Silverado Canyon, V-10-64, on Salvia sp. (crowns and roots) (D. R. Miller and J. A. Froebe, UCD). Placer Co.: Placer, VII-1937, on ? (F. Clark, CDA). Riverside Co.: Arlington, IX-14-42, on Aucuba japónica (C. E. Ehmann, CDA); Beaumont, I X 1922, on Ambrosia sp. (M. C. Tanquary, CDA); Deep Canyon, VII-26-63, on Hyptis emoryi (E. I. Schlinger, UCD); X-9-63, on ? (E. I. Schlinger and M. E. Irwin, UCD); Gilman Springs, XI-20-42, on Agathaea coelestis, Erica sp. (C. E. Ehmann, CDA); XI-13-40, on Primula obconica (C. R. Tower, CDA); Highgrove, VI-18-58, on Vinca sp. (H. S. Elmer, UCD); Indio, XI-2-31, on ? (L. E. Myers,

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CDA); Mira Loma, X-31-42, on Prunus persica (F. R. Piatt and Kilgore, CD A); Perris, IV-23-36, on ? (J. D. Maple, UCD); 2 miles west Perris, IV-23-36, on ? (J. D. Maple, UCD); Riverside, X-16-42, on Abutilon sp. (C. E. Ehmann, CD A ) ; VII-31-35, on Ambrosia ? (J. D. Maple, UCD); VII-6-16, on Aralia sp. (F. Sharp, CDA); VIII-20-29, on Calendula sp. (A. J. Basinger, UCD); IV-15-35, on Citrus sp. (H. J. Quayle, UCD); III-1935, reared in insectary (J. D. Maple, UCD); 1-20-17, on Geranium sp. (A. F. Swain, UCD); VII-28-21, on Solanum sp. (L. E. Myers, CDA); IX-28-16, on Viburnum sp. (collector ?,• CD A ) ; VIII-24-24, on "sunflower" (J. C. Chamberlin, UCD); VI-1925, on "Australian pink vine" (J. C. Chamberlin, UCD). Sacramento Co.: East Sacramento, III—15—30, on Fuchsia sp. (collector ?, CDA); Sacramento (nursery), IV-23-43, on ? (A. E. Morrison, CDA); X I 9-32, on Fatsia sp. (Bachman, CDA); 111-15-30, on Fuchsia sp. (collector ?, CDA); Sacramento, VIII-6-41, on Aucuba sp. (Milbrath, CDA); I X 9-35, on Bouvardia sp. (G. R. Battelle, CDA) ; XII3-42, on Dimorphotheca sp. (J. B. Steinweden, CDA; VIII-7-42, on Fuchsia sp. (H. L. McKenzie, CDA); IV-5-30, on Hibiscus sp. (Bachman, CDA); VI-14-40, on Pelargonium sp. (G. Stout, CDA); IX-28-34, on "rose of Sharon" (M. R. Harris, CDA); X-5-36, on "rose of Sharon" (S. L. Lockwood, CDA). San Benito Co.: Hollister, IX-1-31, on Althaea sp. (H. E. Burke, UCD). San Bernardino Co.: Lost Palm Canyon, IV-8-39, on Chilopsis linearis (R. C. Dickson, UCD); Ontario, VI-15-31, on Asclepias sp. (G. A. Pohl, CDA); III—12—36, on Cytisus hillieri, Viburnum macrocephalum, Peteria sp. (from England) (G. A. Pohl, CDA); VI-15-27, on Pueraria sp. (G. P. Weldon, UCD); III-12-36, on Styrax wilsoni and Elsholtzia stauntonii (from England) (G. A. Prole, CDA); Redlands, X-1922, on "ornamentals" (G. F. Ferris, UCD); San Bernardino, V-1923, on Geranium sp. (J. P. Coy, CDA); XI-1929, on Lantana sp. (J. P. Coy, CDA); Uplands, XII-1920, on Geranium sp. (H. S. Smith, UCD). San Diego Co.: 1 mile east Boulevard, VIII10-64, on Gutierrezia microcephala (J. A. Powell and J. A. Chemsak, UCD); Chula Vista, V-6-63, on Jasminum sp. (W. C. Reed, CDA); (Chula Vista quarantined at San Ysidro) XII-6-46, on Saintpaulia sp. (F. E. Swan, USNM); San Diego, IV-6-56, on Aralia sp. (G. S. Hill, CDA); XII-31-28, on Lantana sp. (R. R. McLean, CDA); IX-28-28, on Rudbeckia sp. (R. R. McLean, CDA); Oceanside, VIII-23-64, on Haplopappus venutus vernonioides (D. R. Estes, UCD); San Ysidro, X-6-40 and X - l 8 - 4 0 , on Geranium sp. (F. E. Swan, USNM); 2 miles south Valley Center, IH-27-64, on Compositae (associated with ant, Formica pilicornis (D. R. Miller and J. A. Froebe, UCD). San Francisco Co.: San Francisco (quarantined at Butte Co.), 1-27-34, on Fuchsia sp. (collector ?, (quarantined at Managua, Nicaragua) X-19-59, on

"ornamental plant" (Estrada, USNM); (quarantined at San Ysidro, San Diego Co.) IX-13-40, on Lantana sp. (F. E. Swan, USNM). San Joaquin Co.: Escalon, XI-7-27, on ? (L. E. Myers, LEM); Tracy, IV-28-33, on Geranium sp. (A. R. Tugel, CDA). San Mateo Co.: Belmont, XII-10-52, on Geranium sp. (E. L. Labadie Jr., CDA); San Mateo, VII-1-5 3, on Sparmannia africana (C. W. Bridges, CDA). Santa Barbara Co.: Carpinteria, X - l 7 - 2 8 , on Veronica sp. (H. C. Lewis, CDA); Santa Barbara, date ?, on Lantana sp. (F. C. Greer, UCD). Santa Clara Co.: Palo Alto, IX-11-44, on Phaseolus sp. (G. F. Ferris, UCD); San Jose, IX-14-44, on Lotus scoparius (J. B. Steinweden, CDA); I V 28-25, on Medicago sativa (L. E. Myers, LEM); Stanford University, X-l6-34, on Helianthus tuberosus (collector ?, UCD); IX-1945, on Hibiscus sp. (G. F. Ferris, UCD); Sunnyvale (nursery), V I 16-52, on Philodendron sp. (G. A. Prole, CDA). Santa Cruz Co.: Green Valley, X - 2 5 ^ 6 , on Fuchsia sp. (T. B. Gallion and R. W. Cogswell, CDA); Watsonville, IX-24-46, on Fuchsia sp. (R. W. Cogswell, CDA); IX-1944, on Lotus scoparius (J. B. Steinweden, CDA). Solano Co.: Suisun, XI-8-43, on Acacia sp. (S. V. Weimar, CDA); Vallejo, XI-12-43, on Helianthus, sp. (S. V. Weimar and G. G. Anderson, CDA); IX-20-39, on Senecio mikanioides (E. O. Essig, CAS). Sonoma Co.: Locality ?, XI-20-29, on Chrysanthemum sp. (Bremner, CDA). Stanislaus Co.: Salida, I X - 2 2 ^ 7 , on "large malvaceous shrubs" (R. W. Harper and R. P. Allen, CDA). Tehama Co.: Corning, X-l0-28, on Pyracantha sp. (E. L. Smith, CDA); Red Bluff, IX-16-41, on Hibiscus sp. (G. P. Merrill, CAS). Tulare Co.: Exeter, X-4-62, on Lantana sp. (Clark, CDA); IX-26-39, on Solanum pseudocapsicum (F. G. Lackland, CDA); Lindsay, 1923, on "ornamentals" (A. J. Filbert, CDA); X - l 8 - 2 8 , on Zantedeschia sp. (A. E. McGregor, UCD). Tuolumne Co.: Sonora, XI-22-61, on Chrysanthemum sp. (H. Hinkley, UCD). Ventura Co.: Camarillo, V-22-59, on Datura sp. (E. I. Schlinger, UCD); Oxnard, IX-28-43, on Asparagus sp. (W. M. Dunning, CDA); Fillmore, V-2-35, on Marrubium vulgare (J. D. Maple, UCD); near Piru, VI-1924, on Carya sp. (E. T. Todds, UCD); Santa Paula, VIII-18-43, on Lycopersicon sp. (J. L. Schall, CDA); 1-1923, on Urtica sp. (H. S. Smith, UCD). Yolo Co.: Davis, X-l937, on Calendula sp. (J. Skoss, CAS); X-3-58, on Chenopodium sp., Datura sp., Marrubium sp., and Solanum sp. (W. H. Lange, UCD); IX-7-61, on Encelia farinosa (H. L. McKenzie, UCD); IX-20-62, on Malva sp. (F. C. Raney, UCD); Woodland, X-1938, on Acacia sp. (S. L. Lockwood, CDA); 1-28-43, on Echium sp. (J. B. Steinweden and C. Hardy, CDA).

TAXONOMY

Yuba Co.: Marysville, XI-17-41, on Parthenocissus quinquefolia, "rose of Sharon," etc. (A. Worledge, CDA); 1-18-43, on Pelargonium sp. (A. Worledge, CDA); VII-15-39, on "perennial plant" (H. A. Crane, CDA).

Phenacoccus graminosus McKenzie Ryegrass Mealybug (Figure 99; Color Plates III and XII) Phenacoccus graminosus McKenzie, 1960. Hilgardia 29(15):717-721, illus. Phenacoccus graminosus McKenzie, 1961. Ibid. 31 (2) :30. Phenacoccus graminosus McKenzie, 1964. Ibid. 35(10):245-246. Phenacoccus graminosus McKenzie, De Lotto, 1964. Bull. Brit. Mus. (Nat. Hist.) Ent. 14(8) :367368, illus. Type Locality and Host. Berkeley, Alameda County, California on ryegrass, Lolium sp. (Gramineae). North American Distribution. Known only from California. Additional Hosts. Avena fatua (Gramineae), Bromus rigidus (Gramineae), Festuca ? sp. (Gramineae), grass (Gramineae), Hordeum sp. (Gramineae), Medicago sativa (Leguminosae), Phalaris tuberosa (Gramineae), Trifolium fragiferum, T. repens (Leguminosae) and Triticum sp. (Gramineae). External Features and Habitat. This mealybug is covered with a fluffy white, powdery secretion. Its waxy filaments extend around the body and are com-

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paratively short, broad, and very jagged. The body color is white in young females, gradually turning pink to darker red in the older specimens (see col. pis. Ill and XII). Body contents are yellowish when crushed; legs cream-yellow with joint articulations brownish-red. A very loose, amorphous ovisac is formed ventrally and posterior to female. The eggs are white. The mealybug is usually found clustered along the sheathing leaf base, but also has been observed exposed on the leaves, and occasionally on the crown of its host. Infrequently, a specimen with ovisac will be found between old leaves on the ground. Recognition Characters. Adult female, mounted, 2.25 to 3.40 mm long, 1.10 to 1.90 mm wide; body form broadly oval; anal lobes fairly well developed. Dorsum normally with 18 pairs of cerarii. Anal lobe cerarius with 2 conical setae, 3 or 4 smaller auxiliary setae, slight concentration trilocular pores. Cerarii on prothorax and head, for the most part, represented by 3 small, conical setae, limited number scattered trilocular pores; remaining cerarii with but 2 conical setae, scarcely any concentration trilocular pores; all cerarii, except those of anal lobes, without auxiliary setae. Dorsal multilocular disk pores present in considerable numbers on all abdominal segments except the last, others distributed, mostly in groups, submarginally on thorax and head. Minute circular pores sparsely scattered. Trilocular pores rather abundant, evenly beset over entire dorsum. Tubular ducts with slight oral collar sparsely distributed over dorsum. Body setae sparse, all small and slender. Anal ring of normal form and size for genus; its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores numerous on all abdominal segments, extending across segment to lateral margins, except on segments 2 and 3 where they occur as a submarginal group; a variable number also present on thorax between spiracles and body margin. Quinquelocular pores considerably smaller than multilocular pores, rather abundantly distributed in median region from fifth abdominal segment forward onto thorax, few on head. Minute circular pores sparsely scattered. Trilocular pores rather evenly distributed on venter except in sternal area. Oral-collar tubular ducts with slight oral rim numerous on last 5 or 6 abdominal segments, sparsely scattered along submargin of thorax. Body setae small and slender, some noticeably longer than those on dorsum. Circulus present, slightly laterally oval, not divided by segmental line, situated about in middle of segment 4. Legs large and comparatively long; claw with prominent denticle or tooth on plantar surface. Antennae 9-segmented, moderately long. Mouthparts broad. Notes. This species is rather closely related to Phenacoccus eremicus Ferris and P. injernalis McKenzie, but differs in possessing dorsal multilocular disk pores on thorax, and ventrally in submarginal groups along thorax. In the two species last men-

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Fig. 99. Phenacoccus graminosus McKenzie, collected on ryegrass, Lolium sp. (Gramineae), at Berkeley, Alameda County, California.

COLOR PLATE XII. The ryegrass mealybug, Phenacoccus graminosus McKenzie, infesting "ripgut" grass, Bromus rigidus (Gramineae), Olivehurst, Yuba County, California.

tioned, these structures are lacking on both surfaces of the thorax. Dr. Harold Morrison examined specimens of P. graminosus before it was described, and in correspondence to me, dated August 12, 1958, he made the following remarks: "This seems as you indicate to be clearly separated from anything else thus far reported from the United States. There are several Phenacoccus species described from other parts of the world that have dorsal multiloculars, and some of these have been reported from grass, but so far as can be told from the descriptions none of these has a ventral circulus. Your report of collection on several hosts in the Bay Area would tend to make one suspicious of possible introduction." DeLotto (1964) records P. graminosus from Africa, and recently the species was collected in New Zealand. These records, along with its previously known presence in Australia, tends to support the statements made in the foregoing paragraph. California Records (Map 90) Alameda Co.: Alameda, VIII-20-53, on "ryegrass" (W. A. Kroger, CD A ) ; Berkeley, IX-24-53, on Lolium sp. (R. P. Allen and R. T. Straw, CD A ) ; Hayward, IX-25-53, on Lolium sp. (R. P. Allen, CDA); Oakland, IX-23-53, on Hordeum sp. (R. P. Allen and J. V. Lonergon, CDA); IX-17-53, on Lolium sp. (J. V. Lonergon and Ritz, CDA); IX- 22-53, on Lolium sp. (R. P. Allen and R. T. Straw, CDA); IX- 11-53, on "ryegrass" (J. V. Lonergon, CDA); Pleasanton, IX-25-62, on "grass" (T. R. Haig, CDA); San Leandro, IX-23-53, on "bromegrass" (S. W. Sibray, CDA). Butte Co.: Chico, IX-24-62, on Festuca ? sp. (T. R. Haig, CDA); 111-29-65, on Bromus rigidus (T. R. Haig and W. W. Wiard, CDA). Contra Costa Co.: Mt. Diablo, IV-5-64, on Gramineae, "in soil and under rocks" (D. R. Miller and J. A. Froebe, UCD); San Pablo, X-10-53, on "ryegrass" (D. J. Bingham, CDA). Humboldt Co.: Areata, VIII-5-64, "under boards" (J. Spallino and T. R. Haig, CDA); 0.5 mile south Bridgeville, VI-13-63, on "leaf mold including wild oats" (W. W. Stanger, UCD). Marin Co.: Greenbrae, IV-2-60, on Avena fatua (T. R. Haig, CDA); San Rafael, X-10-53, on "ryegrass" (D. J. Bingham, CDA). Monterey Co.: North Salinas, 111-21-55, on Hordeum sp. (R. P. Allen and R. V. Emparan, CDA). Napa Co.: Napa, 1-15-59, on Lolium sp. (crown) (H. L. McKenzie and R. P. Allen, UCD). Santa Clara Co.: Sunnyvale, IV-7-59, on "lawngrass and grass" (T. R. Haig and E. Winkler, CDA). Santa Cruz Co.: Santa Cruz, VI-19-56, on grass (L. R. Gillogly, CDA); 4 miles north Watsonville, VI-25-54, on Lolium sp. (R. P. Allen and L. R. Gillogly, CDA). Solano Co.: Vallejo, XII-21-56, on "grass" (under walnut tree) (D. R. Haug, CDA). Yolo Co.: Olivehurst, IH-24-65, on Avena fatua (T. R. Haig, CDA); 7 mile west Winters, IV-16-66, on "grass" under rock (R. F. Wilkey, CDA).

Phenacoccus helianthi (Cockerell) Sunflower Mealybug (Figure 100) Pseudococcus helianthi Cockerell, 1893. Entomologist 26:352. Phenacoccus helianthi (Cockerell), Cockerell, 1894. Canad. Ent. 26:285. Phenacoccus helianthi (Cockerell), Cockerell, 1900. Entomologist 33:86. Phenacoccus helianthi (Cockerell), Ferris, 1919. Stanford Univ. Pubs., Univ. Ser., p. 22. Phenacoccus helianthi (Cockerell), Ferris, 1921. Stanford Univ. Pubs. Univ. Ser., Biol. Sci. 1(2):84-85. Paroudablishelianthi (Cockerell), Borchsenius, 1949. Akad. Nauk. Zool. Inst., n.s. 38(7) :238. (In Russian). Phenacoccus helianthi (Cockerell), Ferris, 1950. Atlas of scale insects of North America, Ser. V: 148-149, illus. Phenacoccus helianthi (Cockerell), McKenzie, 1961. Hilgardia 31(2):26. Type Locality and Host. Las Cruces, Dona Ana County, New Mexico, on sunflower, Helianthus sp. (Compositae). North American Distribution. Arizona, California, Colorado, Mexico, New Mexico, and Texas. Additional Hosts. Baccharis ? sp. (Compositae), Bertholletia sp. (Lecythidaceae), Bouteloua sp. (Gramineae), Bursera microphylla (Burseraceae), Cassia sp. (Leguminosae), Ceiosia floribunda (Amaranthaceae), Citrus sinensis (Rutaceae), Codiaeum sp. (Euphorbiaceae), Grass (Gramineae), Franseria dumosa (Compositae), Gutierrezia sp. (Compositae), Haplopappus sp. (Compositae), Hymenoclea salsola

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Fig. 100. Phenacoccus helianthi (Cockerell), collected on Hymenoclea salsola (Compositae), at Townes Pass, Inyo County, California. (Illustration prepared from specimens collected on Pluchea sp. [Compositae], Mesilla Park, Dona Ana County, New Mexico.)

TAXONOMY (Compositae), Kallstroemia sp. (Zygophyllaceae), Larrea sp. (Zygophyllaceae), Parosela spinosa (Leguminosae), Pinus ponderosa (Pinaceae), Pluchea sericea, Pluchea sp. (Compositae), (Polemoniaceae), Prosopis sp. (Leguminosae), Ptiloria sp. (Compositae), Salvia carnosa (Labiatae), Solatium douglasii, Solanum sp. (Solanaceae), and Stephanomeria sp. prob. virgata (Compositae). External Features and Habitat. The body of this species is thinly covered with a waxy secretion, the lateral filaments are perhaps half the width of the body, and the caudal filaments are almost as long as the body. A slender ovisac is formed—this reaching a length of as much as 10.00 mm. The mealybug occurs on the leaves and stems of its host. Recognition Characters. Adult female, mounted, 2.20 to 4.20 mm long, 1.50 to 2.20 mm wide; body form broadly oval. Dorsum normally with 18 pairs of cerarii. Anal lobe cerarius with from 6-15 moderately large, somewhat lanceolate setae, slight concentration of trilocular pores, borne on distinct sclerotized area. Penultimate cerarius with slightly smaller group of conical setae, somewhat less distinct sclerotized area; antepenultimate cerarius with 4-5 conical setae, no sclerotization; remaining cerarii with usually only 2 conical setae except in head region where there may be 3-4, these somewhat smaller than those of anal lobes. Multilocular disk pores on dorsum in a row along posterior margin of each abdominal segment, scattered over thoracic region. Trilocular pores numerous, evenly beset over entire dorsum. Minute circular pores scattered. Small tubular ducts of oral-collar type in considerable numbers on dorsum, scattered over thoracic region. Body setae very small, of same shape as those of cerarii, sparsely distributed over entire dorsum. Anal ring apical, with no unusual features; each of its 6 setae about twice as long as greatest diameter of ring. Venter with numerous multilocular disk pores on all abdominal segments forward to fourth and in lateral areas of some of these segments. Quinquelocular pores moderately numerous in midregion from anterior segments of abdomen forward to head. Trilocular pores rather evenly distributed, except in sternal area of thorax. Minute circular pores scattered. Tubular ducts of same size and form as on dorsum present on abdomen, especially so in lateral regions, also distributed along submargins and in midregion of thorax. Body setae slender, some longer than those on dorsum. Circulus oval, ends slightly produced laterally and usually sclerotized. Legs comparatively large and well developed; hind tibia with a few translucent pores at distal end; claw with prominent denticle on plantar surface. Antennae normally 9-segmented, moderately long. Notes. This species is quite closely related to Phenacoccus cajonensis McKenzie, a species described as new in this study, but differs principally in possessing ventral quiquelocular pores especially on anterior abdominal segments and in sternal region;

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penultimate cerarius sclerotized with a cluster of from 5-12 setae; and ventral multilocular disk pores absent on thorax. On the other hand, P. cajonensis rarely has ventral quinquelocular pores around mouthparts; penultimate cerarius is unsclerotized with usually 2, rarely 3, setae; and possesses numerous ventral submarginal clusters of multilocular disk pores on thorax. In addition, the species last mentioned appears to be quite specific on Eriogonum spp. (Polygonaceae), whereas P. helianthi seems to be a more general feeder. California Records (Map 91)

Imperial Co.: Calexico, 11-28-57, on Pluchea sericea (G. G. Beevor, CDA); near El Centro, 1-3064, on Prosopis sp. (A. L. Morrison, CDA); 10 miles west Ocotillo, IV-7-63, on Parosela spinosa (R. L. Westcott, UCD) ; Seeley, VIII-21-56, on "mesquite" ? (H. H. Blakemore, CDA) ; 1-24-57, on Pluchea sericea (G. G. Beevor, CDA). Inyo Co.: Darwin Falls, VII-16-46, on Compositae (G. F. Ferris, UCD); Little Lake, VI-24-64, on Franseria dumosa and Haplopappus sp. (foliage) (D. R. Miller, UCD) ; Townes Pass, VI-12-63, on Hymenoclea salsola (foliage) (D. R. Miller, UCD). Los Angeles Co.: Lancaster, VI-12-18, on Ptiloria sp. (G. F. Ferris, UCD); VIII-1919, on Ceiosia ftoribunda (G. F. Ferris, UCD); Los Angeles, IX6-1892, on Citrus sinensis (D. W. Coquillett, USNM). Riverside Co.: 2 miles south Coachella, VI-10-63, on Baccharis ? sp. and Pluchea sericea (foliage) (D. R. Miller, UCD); Prado Dam, VII-24-63, on Stephanomeria (probably virgata) (foliage) D. R. Miller,

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UCD); Riverside, IX-8-1892, on Solanum douglasii (D. W. Coquillett, USNM) ; IX-8-1892, on Solanum sp. (Claflin, USNM); 1 mile west Travertine Rock, Hwy. 99, VIII-6-63, on Hymenoclea salsola (W. H. Ewart, UCD). San Bernardino Co.: Apple Valley (near Victorville), V-16-48, on Larrea sp. (W. D. Dyer, CDA) ; Cajon Pass, IV-9-60, on Salvia carnosa (W. H. Ewart, UCD); Morongo Valley, VI-9-63, on Baccharis ? sp. (foliage) (D. R. Miller, UCD).

Phenacoccus infernalis McKenzie Infernal Mealybug (Figure 101) Phenacoccus infernalis McKenzie, 1962. Hilgardia 32(14):652-654, illus. Phenacoccus infernalis McKenzie, 1964. Ibid. 35 (10):250. Type Locality and Host. Tombstone (Boothill Cemetery, Cochise County, Arizona, on ocotillo, Fouquieria splendens (Fouquieriaceae). North American Distribution. Arizona and California. Additional Hosts. None. External Features and Habitat. According to the collectors, this species was found feeding on the leaves and stems of its host. No other information available at this time. Recognition Characters. Adult female, mounted, 2.25 to 3.75 mm long, 1.25 to 2.00 mm wide; body form broadly oval. Dorsum normally with 18 pairs of cerarii. Anal lobe cerarius with 2 conical setae, 3 or 4 smaller setae, slight concentration of trilocular pores. Remaining cerarii with but 2 cerarian setae, these generally quite small, without auxiliary slender setae, slight concentration of trilocular pores, often borne on more or less distinct prominences particularly in early adult specimens. Dorsal multilocular disk pores present in considerable numbers of all abdominal segments except posterior apical, second, and intermediate area of third, absent on thorax and head. Trilocular pores rather sparse, evenly beset over entire dorsum, each pore appearing with numerous irregular lines. Oral-collar tubular ducts sparsely distributed on abdomen, absent on thorax and head. Body setae small and slender, sparse. Anal ring of normal form and size for genus; each of its 6 setae about twice as long as greatest diameter of ring. Venter with considerable numbers of multilocular disk pores on all abdominal segments except second, extending across each segment to lateral margins, except on segment 3 where 1 or 2 pores may occur submarginally; absent on thorax and head. Quinquelocular pores occur in small numbers near mouthparts. Trilocular pores rather evenly distributed. Small, rather elongate tubular ducts with slight oral rim situated in considerable numbers on last 5 or 6 abdominal segments, sparsely distributed anterior to

these. Body setae slender, some considerably longer than those on dorsum. Circulus quite large, normally appearing broadly oval, capable of folding along intersegmental line. Legs comparatively large and long; apical half of hind tibiae sometimes with few small translucent pores; claw with prominent denticle or tooth on plantar surface. Antennae normally 9-segmented, moderately long. Notes. This mealybug is rather closely related to Phenacoccus alleni McKenzie, but differs principally in the absence of oral-collar tubular ducts on dorsal and ventral surfaces of thorax; and in its apparent preference for a single host, "ocotillo," Fouquieria splendens (Fouquieriaceae). On the other hand, P. alleni, possesses oral-collar tubular ducts on both dorsal and ventral surfaces of thorax; and occurs on numerous hosts but, to date, has never been taken on Fouquieria splendens. Phenacoccus infernalis was originally described from Arizona. During 1963, specimens of this mealybug were found on the type host, ocotillo, in a nursery at Conoga Park, Los Angeles County, California. The infested plants originated in Imperial County, according to the peddler who sold them to the nursery. California Records (Map 92) Los Angeles Co.: Conoga Park, IV-18-63, on Fouquieria splendens (D. Williams, CDA).

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Fig. 101. Phenacoccus infernalis McKenzie, collected on ocotillo, Fouquieria splendens (Fouquieriaceae), at Tombstone (Boothill Cemetery), Cochise County, Arizona.

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Phenacoccus lotearum McKenzie Lotus Mealybug (Figure 102) Phenacoccus lotearum McKenzie, 1960. Hilgardia 2 9 ( 1 5 ) : 720-723, illus. Phenacoccus lotearum McKenzie, 1964. Ibid. 35 (10):250. Type Locality and Host. Summit, Santa Susana Pass, Ventura County, California, on bird's-foot trefoil, Lotus sp. (Leguminosae). North American Distribution. Known only from California. Additional Hosts. Anaphalis margaritacea (Compositae), Centaurea melitensis (Compositae), Chaenactis tenuifolia (Compositae), Cynodon dactylon (Gramineae), Eriogonum sp. (Polygonaceae), and Stephanomeria sp. (Compositae). External Features and Habitat. This mealybug is grayish, covered with a heavy, fluffy, wax secretion, has very inconspicuous caudal filaments and reddish legs. The body contents of the insect are gray when crushed. Ovisacs were not observed. The species occurs on the crown and roots of its host, and has been collected in close association with ants, Crematogaster coarctata vermiculata and Solenopsis truncorum. The ants were observed fondling and carrying the mealybugs. Recognition Characters. Adult female, mounted, 2.40 to 4.40 mm long, 1.90 to 2.50 mm wide; body form broadly oval to rotund. Dorsum with cerarian pairs reduced, normally only 4 or 5 recognizable counting forward from anal lobes. Anal lobe cerarius with 2 rather small, slender, conical setae, few, usually 3, slender auxiliary setae, limited number of scattered trilocular pores. Remaining cerarii with 2 conical setae, these, from anal lobes forward, progressively small, more slender, and usually more widely separated than anal lobe pair, no auxiliary setae, and scarcely any concentration of trilocular pores. Trilocular pores seem somewhat larger than usual, sparsely beset over entire dorsum, each pore appearing irregularly broken with numerous fine lines. Tubular ducts absent, except for an enlarged submarginal oral-collar duct either anterior to, or within confines of, anal lobe cerarius. Dorsal body setae sparse, all small and slender. Anal ring apical, with no unusual characters; each of its 6 setae about twice as long as greatest diameter of ring. Venter with variable limited numbers of multilocular disk pores situated on last abdominal segment forward to posterior margin of fifth. Quinquelocular pores absent. Trilocular pores seem somewhat smaller than those on dorsum, evenly distributed throughout entire venter. Small tubular ducts of oral-collar type occur sparsely in same general area on abdomen as multiloculars, absent elsewhere on venter. Body setae slender, some noticeably longer than those of dorsum. Circulus absent. Legs large, well developed, femora and tibiae robust; tibiae with 35 to 51 translucent pores scattered along entire segment; claw with pro-

minent denticle on plantar surface. Antennae moderately long, either 6-, 7-, or 8-segmented. Notes. This species seems quite closely related to Phenacoccus pauperatus Ferris, but differs in possessing far more numerous ventral multilocular disk pores on abdomen; 4 or 5, usually 4, discernible dorsal abdominal pairs of cerarii; antennae 7- to 8segmented; and legs more robust. P. pauperatus, on the other hand, seldom has more than 10 ventral multiloculars on abdomen, these lying immediately anterior to the vulva; 18 pairs of discernible cerarii on dorsum; 9-segmented antennae; and fairly slender legs. California Records (Map 9 3 )

Los Angeles Co.: 3 miles north Castaic, VI-19-64, on Lotus sp. (crown and roots) (D. R. Miller, UCD); Irwindale (San Gabriel River), H - 1 6 - 6 5 , on Cynodon dactylon, tended by ants, Solenopsis truncorum (R. R. Snelling, U C D ) ; Whittier, III10-35, on Lotus sp. (J. D. Maple, UCD). Orange Co. : 2 miles north Modjeska Canyon, VI7 - 6 3 , on Centaurea melitensis (roots) (D. R. Miller, UCD) ; 9 miles east San Juan Capistrano, VI-22-64, on Lotus sp. (crown) (D. R. Miller, UCD). Riverside Co.: 5 miles southwest Lake Elsinore, VI-21-64, on Lotus sp. (crown) (D. R. Miller, UCD.) San Diego Co. : Bonsall, X I I - 2 9 - 6 4 , on Eriogonum sp. (crown and roots) (D. R. and E. R. Miller, UCD) ; 2 miles east Bonsall, VI-20-64, on Chaenactis tenuifolia (D. R. Miller, U C D ) ; Escondido, VII3-62, on Lotus sp. (R. O. Baker, CDA) ; VIII-2-62, on Lotus sp. (C. A. Avery, C D A ) ; 5 miles east Escondido, X I I - 2 7 - 6 3 , on Lotus sp. (crown and

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Fig. 102. Phenacoccus lotearum McKenzie, collected on bird's-foot trefoil, Lotus sp. (Leguminosae), at Summit, Santa Susana Pass, Ventura County, California.

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roots) (D. R. Miller and J. A. Froebe, UCD); 2 miles south Pala, VI-20-64, on Anaphalis margaritacea (roots) (D. R. Miler, UCD); 1 mile northwest Valley Center, VI-23-64, on Stephanomeria sp. (roots) (D. R. Miller, UCD). Ventura Co.: Fillmore, II-8-36, on Lotus sp. (J. D. Maple, UCD); Summit, Santa Susana Pass, on Lotus sp. (J. D. Maple, UCD).

Phenacoccus lycii (Ferris) Desert Thorn Mealybug (Figure 103) Pseudococcus lycii Ferris, 1919. Stanford Univ. Pubs., Univ. Ser., pp. 30-31. Phenacoccus lycii (Ferris), Ferris, 1950. Atlas of scale insects of North America, Ser. V:150151, illus. Type Locality and Host. Cottonwood, Yavapai County, Arizona, on desert thorn, Lycium sp. (Solanaceae). North American Distribution. Arizona and California. Additional Hosts. Found only on Lycium sp. (Solonaceae). External Features and Habitat. The general appearance of this mealybug is very much like that of a soft scale. The body color is yellow, and there are irregular transverse brownish bands. No filaments are present, and a white secretion occurs on the underside of an almost circular body. No ovisac is produced, and there were no ant associations. This mealybug occurs on the roots of its host. Recognition Characters. Adult female, mounted, 2.50 to 4.75 mm long, 2.00 to 4.20 mm wide; body form of fully grown female almost circular, very rotund. Accompanying illustration made from specimens which have just entered last instar and are not yet greatly distended. Dorsum normally with 17 pairs of cerarii, each with but 2 large, conical or slightly lanceolate setae, except for frontal cerarius which may have 2 or 3 setae of different sizes, most setae practically same size throughout, no auxiliary setae, very little if any concentration trilocular pores, no basal sclerotization. Trilocular pores relatively large, sparsely beset on dorsum. Oral-collar tubular type ducts extremely large, few, with sclerotized tubes, reduced or obsolete oral collars, without filamentous inner prolongation that is usually evident in ducts of Pseudococcidae, one situated on each side near anal lobes, remainder totaling less than 20, in thoracic and cephalic regions. Body setae few, mostly of lanceolate form, considerably smaller than those of cerarii. Anal ring of normal form, comparatively large, at full maturity deeply retracted between anal lobes; each of its 6 setae about twice as long as greatest diameter of ring itself. Venter without multilocular disk pores. Quinquelocular pores present in midregion of abdomen from above vulva on eighth abdominal segment forward to fifth segment, apparently absent in thoracic and head regions. Trilocular pores confined to extreme lateral

areas from anal lobes forward to head. Oral-collar tubular ducts of conventional type, smaller than those on dorsum, occur in numbers in lateral areas of abdomen, abundant in sternal and lateral areas of mesothorax, prothorax, scattered on head. Body setae slender, sparse, some noticeably longer than those on dorsum. Circulus basically oval, very faintly defined in fully developed adult females, apparently not divided by intersegmental line. Legs quite stout; tarsus about one-third as long as tibia; claw with well developed, very distinct denticle. Antennae 7- or 8-segmented, division between seventh and eighth segments at times partial. Notes. This species seems related to Phenacoccus destitutus McKenzie, a California species described as new in this book, but differs in the absence of ventral quinquelocular pores on the thorax and head, and in possessing 7- or 8-segmented antennae. In the species last mentioned, on the other hand, ventral quinquelocular pores are present in numbers on the thorax and head, and the antennae are 9-segmented. In addition, Phenacoccus lycii possesses numerous oralcollar tubular ducts in clusters in sternal and lateral areas of the mesothorax and prothorax, whereas these structures are absent in P. destitutus. California Records (Map 94) San Bernardino Co.: Yucca Valley, VI-9-63, on Lycium sp. (roots) (D. R. Miller, UCD).

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Fig. 103. Phenacoccus lycii (Ferris), collected on Lycium sp. (Solanaceae), at Yucca Valley, San Bernardino County, California. (Illustration prepared from type material.)

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Phenacoccus megaulus McKenzie, new species Large-Duct Mealybug (Figure 104) Type Locality and Host. 5 miles east Big Pine, Inyo County, California, on a composite, Stephanomeria sp. prob. pauciftora (Compositae), July 18, 1963, by D. R. Miller. Type Material. Holotype adult female (single specimen on slide) deposited in museum, University of California, Davis. North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. This species has cephalad, lateral, and posterior filaments, all short, except the 2 apical, caudal pairs which are at least twice as long as the others. The body contents are dark green when crushed, although this color does not show through the heavy, white, waxy secretion covering body. The legs are bright red. This mealybug was collected by beating the foliage of its host. It presumably feeds on the foliage and stems of Stephanomeria. Recognition Characters. Single adult female (here considered quite young), mounted, 2.40 mm long, 1.60 mm wide; body form broadly oval. Dorsum normally with 18 pairs of cerarii. Anal lobe cerarius with as many as 16 enlarged, slightly lanceolate, setae, accompanied by as many or more trilocular pores, all set in slightly sclerotized area. Remaining cerarii with 5-9 somewhat smaller setae of same form associated with slight group of trilocular pores, slight sclerotization. Dorsal median group of cerarii, with setae much smaller than those in marginal cerarii, occur in median area of eighth abdominal segment, in submarginal band from fifth abdominal segment forward to prothorax, and in midregion of head. Trilocular pores evenly beset over entire dorsal surface, each pore appearing irregularly broken with fine lines. Minute circular pores scattered over surface. Dorsum with few, relatively very large, tubular ducts, these almost equal in size to outer ring of ventral modified multilocular (quinquelocular?) disk pores, provided with distinct oral collar. These ducts, variable in number, occur mainly in a broken submarginal row from sixth abdominal segment forward to mesothorax, 2 to 4 on eighth segment, 1 or 2 usually scattered in median area of thorax and head. Body setae short and slender. Anal ring apical, with no unusual features; each of its 6 setae about twice as long as greatest diameter of ring. Venter with modified multilocular (quinquelocular?) disk pores present in considerable numbers from median area of fifth abdominal segment forward to prothorax, usually absent on head. Trilocular pores similar to those on dorsum distributed fairly evenly except for certain "clear" areas in sternal region. Minute circular pores scattered over venter. Very few

oral-collar tubular ducts, smaller than those on dorsum, situated on 8-7, rarely forward to fifth, abdominal segments, apparently absent elsewhere on venter. Body setae slender and generally longer than those of dorsum. Circulus present, distinctly oval, situated slightly above intersegmental line, not divided by this line. Legs proportionately quite large; without noticeable translucent pores on hind tibiae; claw with prominent denticle quite near tip. Antennae 9-segmented, noticeably long and slender. Notes. This mealybug is almost a "twin" species of Phenacoccus eriogoni Ferris, but differs in possessing a variable number of relatively large oral-collar tubular ducts on dorsum, as well as numerous ventral modified multilocular (quinquelocular?) disk pores situated in median area of fifth abdominal segment forward to and including prothorax. In P. eriogoni, on the other hand, tubular ducts of any kind are absent on dorsum, as are ventral modified multilocular (quinquelocular?) disk pores in sternal region, except for possibly an occasional such pore in this general area. California Records (Map 95) Inyo Co.: 5 miles east Big Pine, VII-18-63, on Stephanomeria sp. prob. paucifiora (foliage) (D. R. Miller, UCD). Riverside Co. : Boyd Desert Research Center (Coyote Canyon), II—5—63, on Hofmeisteria pluriseta (W. H. Ewart, UCD); Deep Canyon, IV-25-63, on Peucephyllum sp. (E. I. Schlinger, UCD).

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Fig. 104. Phenacoccus megaulus McKenzie, new species, on composite, Stephanomeria sp. probably paucifiora (Compositae), east of Big Pine, Inyo County, California.

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Phenacoccus minimus Tinsley Minimus Mealybug (Figure 105) Phenacoccus minimus Tinsley, 1898. Canad. Ent. 30:223-224, illus. Phenacoccus kuwanae Coleman, 1903. Jour. N. Y. Ent. Soc. 11:62, illus. Phenacoccus kuwanae Coleman, Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., pp. 58-59. Phenacoccus minimus Tinsley, Ferris, 1950. Atlas of scale insects of North America, Ser. V:152153, illus. Type Locality and Host. Fort Collins, Larimer County, Colorado, on silver spruce, Picea pungens (Pinaceae). North American Distribution. California, Canada, Colorado, and New Mexico. Additional Hosts. Picea breweriana (Pinaceae) and Populus tremuloides (Salicaceae). External Features and Habitat. According to Ferris (1950), "The adult female is entirely enclosed within a sac formed of coarse threads. All of these specimens occurred on the needles, and the species is recorded by Tinsley as occurring in this situation. The specimens recorded by Coleman were said to have been found on lichens on the spruce, to which they evidently had migrated for oviposition." Recognition Characters. Adult female, mounted, 1.60 to 2.60 mm long, 1.00 to 1.80 mm wide; body shape broadly oval. Dorsum of abdomen with cerarii recognizable only on 1 or 2 segments preceding anal lobes. Anal lobe cerarius with 2 quite small, slender, slightly lanceolate setae set far apart, variable number of other very small setae, these accompanied by few scattered trilocular pores, in some specimens surrounded by slightly sclerotized area. Cerarii either absent or so reduced as to be unrecognizable along anterior abdominal segments and thorax, but in cephalic region 4 pairs evident, these with 3 or 4 small, slightly lanceolate setae, few trilocular pores, borne on slight prominences. Trilocular pores numerous over entire dorsum. Oral-collar tubular ducts relatively very large, numerous, arranged in a more or less single row on abdominal segments, elsewhere scattered. In diameter these ducts equal, or even exceed, outer ring of ventral multilocular pores and are provided with distinct oral collar. Body setae very small, sparse. Anal ring apical with no unusual features; each of its 6 setae nearly twice as long as greatest diameter of ring. Venter with multilocular disk pores in midregion of abdominal segments forward to posterior margin of third. Quinquelocular pores scattered from first apparent abdominal segment to head. Trilocular pores rather sparse, noticeable "clear" areas showing in sternal region. Tubular ducts of 2 sizes evident, smaller associated somewhat with multilocular disk pores, larger ones like those on dorsum confined to marginal regions of abdomen and thorax. Body setae

sparse, variable in size, some noticeably longer than those on dorsum. Circulus very small, oval or circular. Legs well developed; claw with prominent denticle or tooth on plantar surface. Antennae normally 9-segmented. Notes. This species resembles, at least to some degree, Phenacoccus dicoriae McKenzie, but differs from it mainly in possessing ventral quinquelocular pores on thorax and head, and a 9-segmented antenna. P. dicoriae, on the other hand, lacks ventral quinqueloculars on thorax, and its antenna is 8segmented. The accompanying illustration of P. minimus was made by Professor G. F. Ferris from specimens collected at Pikes Peak, Colorado. California Records (Map 96) Siskiyou Co.: Near summit of Salmon Mountains, about 10 miles west Salmon Forks, VIII-3-01, on Picea breweriana (G. A. Coleman, UCD).

Phenacoccus multisetosus McKenzie, new species Multiple-Setae Mealybug (Figure 106) Type Locality and Host. Type and paratype adult females on roots of Pteryxia petraea (Umbelliferae), at Westguard Pass, Inyo County, California, collected July 18, 1963, by D. R. Miller. Miller collected additional paratype adult females of this species as follows: on crown and roots of Chrysothamnus sp. (Compositae), 1.5 miles north Hallelujah Junction, Lassen County, California, July 6, 1964; and on

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Fig. 105. Phenacoccus minimus Tinsley, collected on Picea breweriana (Pinaceae), near summit of Salmon Mountains, about 10 miles west Salmon Forks, Siskiyou County, California. (Illustration prepared from specimens collected on Picea pungens [Pinaceae], Pikes Peak, El Paso County, Colorado.)

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Fig. 106. Phenacoccus multisetosus McKenzie, new species, collected on Pteryxia petraea (Umbelliferae), at Westguard Pass, Inyo County, California.

TAXONOMY crown of Tetradymia canescens (Compositae), at Vinton, Plumas County, California, July 6, 1964. Type Material. Holotype adult female (single specimen on slide) and female paratypes in museum, University of California, Davis. Paratypes in United States National Museum, Washington, D. C., and in collection of California State Department of Agriculture, Sacramento. North American Distribution. Known only from California. Additional Hosts. Chrysothamnus sp. (Compositae) and Tetradymia canescens (Compositae). External Features and Habitat. This mealybug is covered with a white, waxy secretion, has numerous minute filaments on thorax and abdomen, and a dark red body. The species is oviparous, producing a filamentous elongate ovisac which covers the entire female, eggs, and the early first instar, yellow-orange, nymphs. The mealybug occurs on the roots of its host. Recognition Characters. Adult female, mounted, 2.10 to 3.50 mm long, 1.70 to 2.00 mm wide; body form broadly oval. Dorsum with 18 pairs of cerarii. Anal lobe cerarius with from 7 to 9, usually 8, conical setae, without auxiliary setae, fairly numerous associated trilocular pores, slight degree of basal sclerotization. Penultimate cerarius with 3 - 5 slightly smaller conical setae; cerarii on abdominal segments 7-6, and those on anterior thorax and head, each often with 3 or 4 conical setae, without auxiliary setae, very slight concentration trilocular pores, little or no basal sclerotization. Remaining cerarii on anterior abdominal segments and thorax with usually 2, seldom 3, conical setae, otherwise similar to those previously discussed. Trilocular pores fairly numerous, evenly beset on dorsum. Multilocular disk pores in considerable numbers across last 6 or 7 abdominal segments, none elsewhere on dorsum. Oral-collar tubular ducts rather numerous especially on abdomen, scattered on thorax and head. Body setae short and slender. Anal ring apical, without unusual characters; each of its 6 setae slightly longer than twice greatest diameter of ring itself. Venter with multilocular disk pores fairly numerous on all abdominal segments, except on segment 3, extending across segment to lateral margins, absent elsewhere on venter. Quinquelocular pores in considerable numbers mainly in midregion from seventh abdominal segment forward to head. Trilocular pores distributed rather evenly on venter. Oralcollar tubular ducts not especially abundant, situated over entire venter, especially noticeable along submargin of body. Body setae slender and generally longer than those of dorsum. Circulus comparatively small, transversely oval, with extraordinary sclerotized processes or spurs arising on each side, apparently not divided by segmental line. Legs comparatively large; claw with prominent denticle on plantar surface near tip. Antennae 9-segmented, fairly long. Mouthparts comparatively broad.

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Notes. This species is related to Phenacoccus gossypii Townsend and Cockerell, but differs in having numerous, usually 8, conical setae, no auxiliary setae in anal lobe cerarius, and a comparatively small, transversely oval circulus with a sclerotized process or spur arising on each side. Phenacoccus gossypii, on the other hand, usually has 3 conical setae, 1 or 2 smaller auxiliary setae in anal lobe cerarius, and the circulus, while often strongly produced laterally, is without a sclerotized process or spur arising on each side. California Records (Map 9 7 ) Inyo Co.: Westguard Pass, V I I - 1 8 - 6 3 , on Pteryxia petraea (roots) (D. R. Miller, UCD). Lassen Co.: 1.5 miles north Hallelujah Junction, V I I - 6 - 6 4 , on Chrysothamnus sp. (crown and roots) (D. R. Miller, UCD). Plumas Co.: Vinton, V I I - 6 - 6 4 , on Tetradymia canescens (crown) (D. R. Miller, UCD).

Phenacoccus pauperatus Ferris Pauper Mealybug (Figure 107) Phenacoccus pauperatus Ferris, 1950. Atlas of scale insects of North America, Ser. V : 154-155, illus. Phenacoccus pauperatus Ferris, McKenzie, 1960. Hilgardia 2 9 ( 1 5 ) : 7 2 3 . Type Locality and Host. Riverside, Riverside County, California, on Eriogonum sp. (Polygonaceae). North American Distribution. Known only from California. Additional Hosts. Eriophyllum confertifiorum (Compositae).

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Fig. 107. Phenacoccus pauperatus Ferris, collected on Eriogonum sp. (Polygonaceae), at Riverside, Riverside County, California.

External Features and Habitat. No available information on external features of this mealybug. The species presumably infests the roots of its host. Recognition Characters. Adult female, mounted, 2.00 to 3.80 mm long, 1.40 to 2.40 mm wide; body shape moderately rotund. Dorsum normally with 18 pairs of cerarii. Anal lobe cerarius with 2 moderately large and 1 or more smaller setae, accompanied by very slight concentration trilocular pores, not borne in a sclerotized area. Remaining cerarii for most part with but 2 setae, these somewhat smaller than those of anal lobes, with slight concentration trilocular pores. Trilocular pores-sparsely beset on dorsal surface, those slightly larger than usual for group, each pore appearing irregularly broken with numerous fine lines. Very few minute circular pores on posterior abdominal segments. Dorsum without multilocular disk pores and apparently destitute of tubular ducts. Body setae very few and very small; eighth abdominal segment with a median dorsal cerarius containing 2 enlarged setae and a few trilocular pores. Anal ring apical; each of its 6 setae about twice as long as greatest diameter of ring. Venter with scarcely more than 10 multilocular disk pores, these usually situated immediately anterior to vulva, infrequently an occasional pore present on seventh and ninth abdominal segments, at times completely absent on abdomen. Trilocular pores sparsely beset throughout, these somewhat smaller than those on dorsum, apparently not internally broken into fine irregular lines. Minute circular pores lightly scattered on ventral surface. Tubular ducts with a slight oral collar sparsely situated near lateral margins of abdominal segments, absent elsewhere on venter. Body setae slender, some noticeably longer than those of dorsum. Circulus absent. Legs well developed; hind tibia with few translucent pores especially near distal end; claw with prominent denticle on plantar surface. Antennae 9-segmented, quite slender. Notes. This species is related to Phenacoccus lotearum McKenzie, but differs from it in possessing only a few ventral multilocular disk pores on abdomen, these lying usually anterior to vulva; 18 pairs of cerarii; and a 9-segmented antenna. P. lotearum, on the other hand, has more numerous ventral multilocular disk pores on abdomen, these situated on last to posterior margin of fifth abdominal segment; only 4 or 5, usually 4, discernible dorsal abdominal pairs of cerarii; and a 6- to 8-segmented antenna. California Records (Map 98) Los Angeles Co.: 7 miles north Castaic, VI-19-64, on Eriogonum sp. (D. R. Miller, U C D ) . Riverside Co.: Riverside, X-23-33, on Eriogonum sp. (S. E. Flanders, U C D ) ; IV-1935, on Eriogonum sp. (J. D. Maple, U C D ) . San Bernardino Co.: Cajon Pass, IV-15-60, in soil under juniper (U. Nur, U C D ) . Santa Clara Co.: Permanente Creek, date ?, on Eriophyllum confertifiorum (collector ?, U C D ) .

Phenacoccus solarti Ferris Solanum Mealybug (Figure 108; Color Plates I and XIII) Pseudococcus solani Cockerell, Essig, 1914. Calif. State Comn. Hort. Monthly Bull. 3 ( 3 ) : 122, illus. (misidentification). Phenacoccus solani Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., pp. 60-61, illus. Phenacoccus solani Ferris, Zimmerman, 1948. Insects of Hawaii, Univ. Hawaii Press, 5:165, illus. Phenacoccus solani Ferris, Ferris, 1950. Atlas of scale insects of North America, Ser. V : 1 5 8 159, illus. Phenacoccus solani Ferris, Ferris, 1953. Ibid., Vol. VI:403. Ferris (1950) states: "There is a possibility that this species is actually a synonym of Pseudococcus solani (Cockerell), which was described in 1894 from potato in New Mexico, but apparently no types of the latter are in the Cockerell collection in Washington, and no statement can be made concerning it." Type Locality and Host. Stanford University, Santa Clara County, California, on tarweed, Hemizonia rudis (Compositae). North American Distribution. California, Colorado, Florida, Louisiana, New Mexico, North Carolina, Oregon, Puerto Rico, and Texas. Additional Hosts. Specimens here considered to be this species are at hand from a long list of hosts, some of which are recorded as follows: Ambrosia sp. (Compositae), Artemisia absinthium and A. heterophylla (Compositae), Aster adscendens (Compositae) Bouteloua sp. (Gramineae), Castilleia applegateii (Scrophulariaceae), Chenopodium sp. (Chenopodiaceae), Cyperus esculentus (Cyperaceae), Echeveria sp. (Crassulaceae), Eriger on sp.

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Fig. 108. Phenacoccus solani Ferris, collected on Solarium tuberosum (Solanaceae), at Merced, Merced County, California. (Illustration prepared from type material.)

MARY FOUY BE «SON

COLOR PLATE XIII. The solanum mealybug, Phenacoccus solani Ferris, on crown of alkali mallow, Sida hederacea (Malvaceae), Davis, Yolo County, California.

TAXONOMY (Compositae), Eucharis amazonica (Amaryllidaceae), Franseria sp. (Compositae), Helianthus sp. (Compositae), Heterotheca sp. (Compositae), Lantana camara (Verbenaceae), Lycopersicon sp. (Solanaceae), Malva sp. (Malvaceae), Narcissus sp. (Amaryllidaceae) (bulbs), Orobanche sp. (crowns) (Orabanchaceae), Polygonum sp. (Polygonaceae), Portulaca sp. (Portulacaceae), Raphanus sativus (Cruciferae), Sida hederacea (Malvaceae), Solanum tuberosum (Solanaceae), Verbesina encelioides var. exauriculata (Compositae), Vigna sinensis (Leguminosae), and Viola sp. (Violaceae). External Features and Habitat. Available information indicates that in life this mealybug is very thinly dusted with a waxy secretion; the marginal filaments are very small or lacking, caudal filaments short though conspicuous. The ovisac is not known, and the species appears to have living young (viviparous). It occurs commonly on the crown and underground portions of its host (see col. pis. I and XIII). Recognition Characters. Adult female, mounted, 2.25 to 3.20 mm long, 1.30 to 1.90 mm wide; body form broadly oval to slightly rotund. Dorsum normally with 18 pairs of cerarii. Anal lobe cerarius usually with 2 moderately stout, silghtly lanceolate setae, 1 to 3 smaller setae similarly shaped, very slight concentration of trilocular pores. Remaining cerarii with setae somewhat smaller, each cerarius with but 2, except perhaps ocular pair, and with not more than 4-5 associated trilocular pores. Dorsum beset with trilocular pores. Minute circular pores scattered. Tubular ducts absent. Body setae evenly beset on dorsum, somewhat lanceolate, the largest scarcely as large as lateral cerarian setae, most of them smaller. Anal ring apical, with no unusual features; each of its 6 setae about twice as long as greatest diameter of ring. Venter with relatively small number of multilocular disk pores in region of vulva, at times with small numbers of these pores forward to posterior mafgin of fifth segment, sometimes with these pores not present beyond seventh abdominal segment, absent elsewhere on body. Quinquelocular pores absent. Trilocular pores distributed fairly evenly. Minute circular pores scattered. Very few small tubular ducts, with slight oral collar, present in midregion and lateral areas of abdominal segments. Body setae slender, some noticeably longer than those of dorsum. Circulus normally quite small, not divided by intersegmental line; this small size, combined with rather firm surrounding wall, slightly removed from segmental line, suggests that it ordinarily is of definite circular or slightly oval shape. Some specimens have circulus somewhat larger, more lax, and distorted, this probably done in course of preparation. Legs well developed; tibiae usually appearing swollen, translucent pores scattered along entire segment; claw with very slight denticle or tooth on plantar surface. Antennae normally 8-segmented, although eighth segment at times partially divided into 2, at times 9 well developed segments present. At times

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antenna may be 8-segmented on one side of body, 9segmented on other. Some populations appear in which 9-segmented antennae occur frequently, although normal number seems definitely to be 8. Notes. This mealybug is quite widespread throughout California. It is closely related to Phenacoccus dejectus Ferris and P. solenopsis Tinsley. It differs from P. defectus in having the ventral multilocular disk pores situated normally on the last 4 or 5 abdominal segments, rarely restricted to the last 3 abdominal segments; antennae 8-segmented (rarely 9-segmented); and a small, oval, undivided circulus. Phenacoccus defectus, on the other hand, has the ventral multilocular disk pores normally confined to the last 2 abdominal segments; antennae 9-segmented; and usually a larger circulus which is divided by intersegmental line across which it lies. According to Ferris (1950), P. solenopsis "is normally considerably larger, its circulus is usually larger and more lax and becomes involved in the intersegmental fold between the fourth and fifth segments. It is thus susceptible to distortion in preparations and, in addition, is at times somewhat laterally produced. The multilocular pores are normally more numerous and are more concentrated in the region of the vulva, and there is a somewhat more definite concentration of trilocular pores about the anal lobe cerarii." California Records (Map 99) Alameda Co.: (quarantined from Los Angeles to Berkeley), 11-10-58, on "succulent" (N. L. Jones, CDA); Berkeley, XI-10-28, on Calendula sp. (collector ?, CAS); 1-24-49, on Echeveria rubella (G. B. Laing, CDA); 11-20-64, on Solanum sp. (V. B. Whitehead, UCD); Oakland, IX-11-53, on Lolium sp. (J. V. Lonergon, CDA); Pleasanton, IX-25-62, on "grass" (T. R. Haig, CDA); Sunol, IX-3-62, on ? (T. R. Haig and D. A. Taylor, CDA). Alpine Co.: Markleeville, VII-9-64, on Chrysothamnus sp. (inside stems, not in burrows) and Liliaceae (crown) (D. R. Miller, UCD). Butte Co.: Chico, IX-4-62, on Amaranthus sp. (T. R. Haig, CDA); IX-20-34, on Euphorbia maculata (E. E. Fix, CDA); Gridley, IV-30-59, on Ambrosia sp. (A. G. Forbes and R. F. Wilkey, CDA); Oroville, X-6-39, on Rumex sp. (J. F. Fitzgerald, CDA). Colusa Co.: 3 miles north Maxwell, VI-25-63, on Ambrosia psilostachya and Sida hederacea (associated with roots and ants, Tapinoma sessile) (D. R. Miller, UCD). Contra Costa Co.: Antioch, 111-25-65, on Euphorbia sp. (F. Brucata, CDA); Martinez, VIII-1765, on Sida hederacea (R. P. Allen, CDA). Fresno Co.: Fresno, XI-12-57, on Narcissus sp. (bulb) (H. V. Dunnegan, CDA). Glenn Co.: Orland, X-15-32, on Chenopodium sp. (H. S. Smith, UCD); Willows, VI-15-31, on Chenopodium sp. (C. H. Wren, CDA); VIII-5-31, on Hibiscus esculentus (A. C. Browne, CDA). Imperial Co.: Andrade, 111-30-45, on Suaeda sp. (A. J. Hansen and G. G. Anderson, CDA).

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Inyo Co.: Bishop, VII-19-63, on Malvaceae (crowns and roots) (D. R. Miller, UCD). Kern Co.: Arvin, VII-1962, reared on Solatium tuberosum (sprouts) (L. Chiarappa, UCD); Oildale, X-3-57, on Salsola kali (H. L. Wilson, CDA). Los Angeles Co.: Antelope Valley, VII-1925, on Tribulus terrestris (L. E. Myers, LEM); Glendora, VII-23-46, on Salsola kali (M. Miller, LEM); Hynes, V-31-44, on Echeveria sp. (F. Boerner, LEM); La Cañada, X-14-49, on Echeveria sp. (L. E. Myers, L E M ) ; La Puente, XII-8-36, on "weed" (L. E. Myers, LEM); Los Angeles, X-6-33, on ? (Janbert, USNM); Montebello, 1942, on Eucharis amerzonica (L. E. Myers, LEM); Pomona, IV23-26, on Lycopersicon sp. (L. E. Myers, LEM); San Dimas, III—12—63, on Cyperus rotundus (G. O. Poinar, Jr. and D. W. Ricker, CDA); San Fernando, 111-23-54, on Helianthus sp. (Hodge, CDA); Watts, VI-27-63, on Centaurea diluía (T. C. Fuller, CDA). Marin Co.: Larkspur, II—8—55, on Lilium sp. (D. J. Bingham and R. E. Garvey, CDA). Merced Co.: Merced, X-30-39, on Solarium tuberosum (C. H. Kinsley, CDA). Modoc Co.: Cedarville road, 7 miles junction # 3 9 5 , V-21-62, on "composite" (roots) (T. R. Haig, CDA). Nevada Co.: Truckee, VIII-26-65, under rocks (T. R. Haig, CDA). Orange Co.: Anaheim, VIII-8-57, on Arenaria verna (R. Miller and D. H. Byers, CDA); 4 miles east El Toro, VI-7-63, on Ambrosia psilostachya (on stem, associated with Crematogaster coarctata vermiculata) (D. R. Miller, UCD); Garden Grove, VI-18-63, on Plantago sp. (D. H. Byers, CDA); Los Alamitos, XII-29-60, under paper on ground (G. Jackson and J. Johnson, CDA); Newport Beach, VI-6-63, on Abronia sp. (D. R. Miller, UCD); Newport Beach (Back Bay), VII-27-63, on Helianthus annuus (roots) (D. R. Miller, UCD); Between Orange and Olive, X - l l - 5 6 , on "nutgrass" (R. J. Bumgardner, CDA); South Laguna, VII-26-63, on Artemisia californica (roots) (D. R. Miller, UCD); Trabuco Canyon, IV-11-63 and VI-7-63, on Ambrosia psilostachya (D. R. Miller, UCD). Placer Co.: 3 miles west Applegate, VIII-11-64, on Centaurea solstitialis (H. L. McKenzie and D. R. Miller, UCD); VIII-10-64, on Helenium sp., Lotus purshianus, and Plantago lanceolata (roots) (H. L. McKenzie, D. R. Miller, J. A. Froebe, and M. F. Benson, UCD); VIII-8-64, on Penstemon sp. (roots) (D. R. Miller, UCD); Michigan Bluff, X I 5-62, on root under stone (R. R. Snelling, CDA); Roseville, IX-20-42, on "grasses" (range) (S. L. Lockwood, CDA). Plumas Co.: 3 miles east Canyondam, VII-9-64, on Antennaria argentea (crown) (D. R. Miller, UCD); Quincy, VIII-24-62, on Aster adscendens (roots) (T. R. Haig, CDA). Riverside Co.: Arlington, IV-1918, on Solanum tuberosum (A. F. Swain UCD); 3.4 miles northeast Gilman Springs, III-9-63, on Artemisia absinthium (roots) (D. R. MiUer, UCD); Highgrove, XI-9-62,

on Cyperus rotundus (D. W. Ricker, UCD); Palm Springs, VIII-27-42, on Cyperus ? sp. (R. C. Tower, CDA). Sacramento Co.: Isleton, VIII-17-31, on Beta saccharifera (Bachman, CDA); Sacramento, IX-2441, on Heterotheca sp. (crowns) (G. F. Ferris, UCD) ; XI-14-44, on Heterotheca grandiflora (H. L. McKenzie, UCD); X-25-28, on "nutgrass" (roots) (H. H. Keifer, CDA). San Bernardino Co.: Fontana, XI-12-51, on ? (D. C. Lloyd, USNM); 1 mile west Joshua Tree, IV-9-63, on Phacelia sp. (D. R. Miller, UCD); Ontario, VIII-19-59, on Erigeron sp. (G. M. Harper, CDA); VII-17-59, on Verbesina encelioides var. exauriculata and Helianthus annuus (G. M. Harper, CDA); Ontario ?, VIII-1960, on "weed" (roots) (G. M. Harper, CDA); Rialto, VI-15-63, on Tribulus terrestris (roots) (G. M. Harper, CDA); San Bernardino, X-21-52, on "hot house plants" (G. M. Harper, CDA). San Diego Co.: 2 miles east Bonsall, VI-20-64, on Franseria sp. (D. R. Miller, UCD); Carlsbad, X-8-51, on "nutgrass" (L. E. Myers, LEM); Encinitas, VIII-9-56, on "orchid" (D. Sprague, CDA) ; Fallbrook, X-24-34, on "corn" (H. H. Keifer, CDA); Leucadia, V I I I ^ - 5 5 , on Sedum sp. (J. Wilson and D. Sprague, CDA); 0.5 miles north Oceanside, 1-29-64, "under rock" (D. R. Miller and J. A. Froebe, UCD); Ramona, XI-8-56, on Coleus sp. (D. Sprague, CDA); San Diego, IX-16-59, on Euphorbia maculata (C. Beck and F. Varuss, CDA); IX-27-62, on "palm" (J. R. Carlin, CDA); 1923, on Viola sp. (E. O. Essig, UCD); Vista, IX-23-65, on Kalanchoè sp. (Metcalf, CDA). San Francisco Co.: San Francisco, 11-18-50, on ? (A. M. Crull, UCD). San Joaquin Co.: Escalon, 1-12-33, on Aquilegia sp., Malva sp., and Portulaca oleracea (T. Heurlin, CDA); Stockton, IX-29-49, on Chenopodium sp. (F. Hutchings and J. Kalstrom, CDA). San Luis Obispo Co.: 15 miles west New Cuyama, IX-21-65, on Salsola kali (D. W. Ricker, UCD). San Mateo Co.: Belmont, X-14-57, on Cyperus sp. (R. Mickelsen and D. J. Bingham, CDA). Santa Clara Co.: Los Altos, VIII-31-59, on Echeveria sp. (C. Howe, CDA); Palo Alto, IX-28-17, on Solanum tuberosum sativus (roots) (R. W. Doane, UCD); San Jose, V-4-64, on Selaginella sp. (E. C. Winkler, CDA); IX-17-17, on Solanum tuberosum (tuber) (G. F. Ferris, UCD); Stanford University, X-21-16, on Hemizonia rudis (roots) (G. F. Ferris, UCD); VII-7-17, on Raphanus sativus (roots) (G. F. Ferris, UCD). Santa Cruz Co. : Capitola, VII-26-40, on Hippeastrum sp. (F. J. March, CDA). Shasta Co.: 3 miles north Old Station, VI-26-63, on Castilleia sp. (roots, associated with ants, Crematogaster coarctata coarctata) (D. R. Miller, UCD). Stanislaus Co.: La Grange, Xl-18-61, on Rumex sp. (R. P. Allen, CDA); Modesto, XI-13-63, on "cactus" (H. Sinclair, CDA). Sutter Co.: Live Oak, X-5-32, on ? (T. D. Ur-

TAXONOMY

bans, CDA); Yuba City, VIII-26-31 on Althaea sp. (L. S. Jones, CDA). Tehama Co.: Cottonwood, VIII-5-31, on Portulaca oleracea (W. E. Lewis, CDA). Trinity Co.: Burnt Ranch, X-8-62, under rock (with ant) (T. R. Haig, CDA); 9 miles east Burnt Ranch, X-8-61, under rock with ant, Solenopsis molesta validiuscula (T. R. Haig, CDA). Tulare Co.: Exeter, 1-7-64, on "cactus" (Watkins, CDA); Mineral King (8,000'), VIII-26-41, on Artemisia heterophylla (G. F. Ferris, UCD). Ventura Co.: Location ?, VIII-1933, on "water plant" (L. Schall, CDA); Location ?, 1924, on Portulaca sp. (A. H. Call, CDA); Fillmore, H-27-36, on Helianthus sp. (J. D. Maple, UCD); Moorpark, 111-29-63, on Cyperus esculentus (D. W. Ricker, UCD); Santa Paula, date ?, on Malva rotundifolia (E. O. Essig, CAS); XI-8-08, on Solanum xanthi douglasii (P. E. Smith, USNM). Yolo Co.: Davis, V-15-59, on Ambrosia sp. (M. L. Oldham, CDA); X-2-61, on Ambrosia psilostachya (M. E. Irwin, UCD); X-8-62, on Crassulaceae (W. Russell, UCD); IX-23-61, on Poly gonum sp. (roots) (M. E. Irwin, UCD); Davis (Putah Creek), XI-10-60, in "dry soil" (M. E. Irwin, UCD); Knights Landing, VI-24-64, on Sida hederacea (Comfort, CDA). Yuba Co.: Marysville, XI-9-31, on Hedera helix (collector ?, CDA).

Map 99. Distribution in California of Phenacoccus solarli Ferris. Mann -»• San

francitco San Matto Santa Crut

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Phenacoccus solenopsis Tinsley Solenopsis Mealybug (Figure 109) Phenacoccus solenopsis Tinsley, 1898. Canad. Ent. 30:47-48, illus. Phenacoccus solenopsis Tinsley, 1898. Ibid 30:319. Phenacoccus cevalliae Cockerell, 1902. Ibid. 34:325. Phenacoccus solenopsis Tinsley, Fernald, 1903. Mass. Agr. Expt. Sta. Spec. Bull. 88, p. 93. Phenacoccus solenopsis Tinsley, Ferris, 1950. Atlas of scale insects of North America, Ser. V:161163, illus. Type Locality and Host. Mesilla Park (on grounds of Agricultural College), Dona Ana County, New Mexico, on Kallstroemia brachystylos (Zygophyllaceae), and from nests of an ant, Solenopsis geminata. North American Distribution. Arizona, California, Colorado, Mississippi, New Mexico, Washington, D. C., and Texas. Additional Hosts. This species occurs on a long list of hosts, some of which are listed as follows: Achillea sp. (Compositae), Althaea sp. (Malvaceae), Ambrosia artemisifolia (Compositae), Atriplex sp. (Chenopodiaceae), Cevallia sinuata (Compositae), Chenopodiaceae, Compositae (sunflower), Encelia farinosa (Compositae), Enceliopsis sp. (Compositae), Eriophyllum sp. (Compositae), Euphorbia sp. (Euphorbiaceae), Franseria sp. (Compositae), Gossypium sp. (Malvaceae), Helianthus sp. (Compositae), "ink weed" ?, Lupinus sp. (Leguminosae), Orobanche sp. (Orobanchaceae), Solanum sp. (Solanaceae), and Suaeda sp. (Chenopodiaceae). External Features and Habitat. The adult female of this mealybug is quite large, covered with a fine, white, powdery, wax secretion, under which appears a dark-green to almost black body. The marginal wax filaments are only slightly developed, and a quite long (approx. 7.00 mm) fluffy, loose-textured ovisac is formed. This species seems to confine most of its feeding activities to above ground. In certain instances, however, it is found feeding on the crown of its host and a few inches below the soil surface. It also has been discovered in the nest of ants. Recognition Characters. Adult female, mounted, 2.30 to 5.50 mm long, 1.50 to 4.30 mm wide; body form rather rotund. This mealybug is a quite large species, the largest specimens observed attaining a length of approximately 5.50 mm, the majority of specimens, however, are about 4.00 mm long. Dorsum normally with 18 pairs of cerarii. Anal lobe cerarius usually with 2 moderately large, conical setae, 1 or 2 setae of same shape but smaller surrounded by numerous, but not crowded, trilocular pores. Remaining cerarii with but 2 setae, these of same shape only slightly smaller than those of anal lobe cerarii, accompanied by few trilocular pores. Trilocular pores quite evenly distributed on dorsum. Minute circular pores scatterd. Tubular ducts absent. Body setae all smaller than any of the cerarian setae, somewhat lanceolate, few, and scattered.

278

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Fig. 109. Phenacoccus solenopsis Tinsley, collected on Encelia farinosa (Compositae), at Indio, Riverside County, California. (Illustration prepared from specimens collected on Encelia farinosa [Compositae], at Riverside, Riverside County, California.)

TAXONOMY

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Anal ring apical, with no unusual characters; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores quite numerous, these normally situated in compact grouping about vulva, with few or none anterior to posterior margin of seventh segment. Quinquelocular pores absent in any material examined. Trilocular pores distributed fairly evenly. Minute circular porqs scattered. A few very small, oral-collar type ducts scattered over median and lateral areas of abdomen, few situated in lateral and median area of thorax and head. Body setae slender and, for the most part, generally longer than those of dorsum. Circulus usually moderately large, somewhat variable, at times slightly produced laterally, or quite small and oval, often involved in intersegmental fold, rather easily distorted in preparations. Legs well developed; hind tibia with translucent pores along entire segment; claw with very small tooth on plantar surface. Antennae normally 9-segmented. Notes. This species is closely related to Phenacoccus solani Ferris, although a combination of certain characters, including a generally larger size; an antenna normally 9-segmented; a circulus normally larger and more subject to distortion and at times somewhat produced laterally; ventral multilocular disk pores usually more numerous and much more concentrated about vulva; trilocular pores of anal lobe cerarii more numerous and more concentrated, and these pores more numerous over dorsum, differentiate it from the species last mentioned. Phenacoccus solani, on the other hand, is usually a much smaller species; its antenna is more often 8-segmented; its circulus is usually small and oval with rather firm surrounding wall; ventral multilocular disk pores in small numbers in region of vulva forward to posterior margin of segment five; trilocular pores very few on anal lobe cerarii, and these structures are generally less numerous on dorsum. With well-stained specimens available for direct comparison, the two species are easily separated. These differences are difficult to place in a key, and the accompanying key to species of this group is definitely weak in this ara. P. solenopsis also resembles P. defectus but differs in possessing more ventral multilocular disk pores (40 or more) in area surrounding vulva, and with several of these pores on posterior margin of seventh segment, whereas in P. defectus there are usually fewer (20 or less) multiloculars around vulva, and only infrequently a single pore on posterior margin of seventh segment. The accompanying figure of P. solenopsis was made by Professor Ferris from specimens collected on Encelia farinosa, at Riverside, California. California Records (Map 100) Imperial Co.: Brawley, IX-10-62, on A triplex polycarpa (G. M. Harper, CDA); Calexico, 1-11-45, on "weed" (A. J. Hansen and C. G. Anderson, CDA); El Centro, XII-13-19, on "ink weed" (W. M. Davidson, CDA); XI-13-35, on Atriplex sp. (R. F. Wil-

key, CDA); 5 miles north Truckhaven, VI-10-63, on Encelia frutescens and Petalonyx thurberi (foliage) (D. R. Miller, UCD); Winterhaven, X-13-61, on "gourds" (R. Benthey and J. Taylor, CDA); Locality ?, VIII-22-50, on "desert plants" (Harrigan, CDA). Inyo Co.: Surprise Canyon, Panamint Mts. IV22-57, on Enceliopsis sp. (J. I. Stage, CDA). Kern Co.: Salt Dale, near Mojave, IV-26-36, on Suaeda sp. (G. F. Ferris, UCD). Mendocino Co.: Mendocino, VII-6-57, in "ant nest" (J. R. Heifer and R. O. Schuster, UCD). Merced Co.: Planada, X-14-42, on Centromadia pungens (H. Graves, CDA). Orange Co.: Balboa, II-6-36, on ? (J. D. Maple, CDA). Riverside Co.: Blythe, VII-27-47, on Phacelia sp. (J. W. Swain, CDA); 2 miles south Coachella, V I 10-63, on Baccharis ? sp. (foliage) (D. R. Miller, UCD); Indio, XI-2-31, on "alkali-heath," a Chenopodiaceae (L. E. Myers, LEM); Palm Springs, X-21-34, on "Chenopodiaceus plant" (C. Gammon, CDA); IV-1944, on Encelia farinosa (T. D. A. Cockerell, CDA); Riverside, VIII-1-35, on Ambrosia sp. (J. D. Maple, UCD); XII-5-34 and II-8-35, on Encelia farinosa (J. D. Maple, CDA); X-1937, on Stephanomeria sp. (J. D. Maple, UCD). San Bernardino Co.: Cima, IV-1961, on "cactus" (barrel) (W. Cronk, CDA); Twentynine Palms, IV10-62, on Atriplex polycarpa (G. M. Harper, CDA); IV-13-35, on ? (J. D. Maple, UCD); IV-14-35, on Lupinus sp. (J. D. Maple, UCD); Vidal, III-1-54, on ? (R. F. Camblin, CDA). San Diego Co.: San Diego, 1924, on Eriophyllum

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? sp. (R. R. McLean, CD A); VI-15-49, on Euphorbia sp. (roots) (R. W. Harper, F. T. Thorne, and E. T. Gammon, CDA). San Mateo Co.: Half Moon Bay, IX-1937, on Franseria sp. (E. O. Essig, CAS); IX-19-37, on

PLANOCOCCUS Planococcus Ferris, 1950. Atlas of the scale insects of North America, Ser. V: 164. Planococcus Ferris, Ezzat and McConnell, 1956. Md. Agr. Expt. Sta. Bull. A-84 (Tech.), p. 60. Genotype. Pseudococcus citri (Risso). Recognition Characters. Pseudococcidae with moderately oval body form, as mounted. Two pairs of dorsal ostioles present. Typically with 18 pairs of cerarii, there being a preocular pair between frontal and ocular pairs. All cerarii normally with 2 conical setae, except for those in head region, which may have 3-4, without slender auxiliary setae, except perhaps on anal lobes. Anal lobe cerarii usually with but a small number of trilocular pores and without associated sclerotization. Dorsum normally with at most but few tubular ducts, these usually without an oral ring. Anal ring usually well developed, cellular, bearing 6 setae. Venter with multilocular disk pores on abdomen. Circulus present or absent; if present, large, extending across fold between fourth and fifth abdominal segments. Legs well developed; denticle or tooth absent on plantar surface of claw. Ventral side of anal lobes always with slender barlike, sclerotized area extending in from base of anal lobe seta. Notes. The separation of Planococcus from Pseudococcus Westwood is obvious on the basis of the characters given above, but some difficulty has been encountered in limiting the genus in other directions. The presence of 18 pairs of cerarii together with the other indicated characters will, however, usually suffice in recognizing the typical forms. Two species of Planococcus are recorded in North America, and both of these occur in California.

KEY Species of Planococcus in North America: Adult Females (Modified from Ferris, 1950)

Dorsum with several oral-collar tubular ducts situated near each of most abdominal cerarii and scattered across most abdominal segments kraunhiae (Kuwana) Dorsum never with more than a single oral-collar tubular duct near cerarii on last 4 abdominal segments, these structures absent on other abdominal segments citri (Risso)

Franseria bipinnatifida (W. C. Reeves, USNM); Location ?, X - l - 5 3 , on Achillea sp. (E. L. Labadie Jr., CDA). Tulare Co.: Angiola, IV-4-61, on Suaeda fruiticosa (R. P. Allen, CDA).

FERRIS Planococcus citri (Risso) Citrus Mealybug (Figure 110; Color Plates I and XIV) Dorthezia citri Risso, 1813. Mus. d'Hist. Nat. Ann. 20:169-212, 401-431 (415-418), illus. Also issued as Essai sur l'histoire naturelle des oranges (Paris: Dufour, 1813) 74 pp. (58-61). Coccus citri (Risso), Boisduval, 1867. Essai sur l'entomologie horticole (Paris: Donnaud), p. 348. Dactylopius citri (Risso), Signoret, 1875. Ann. Soc. Ent. de France (Ser. 5) 5:312, illus. Lecanium phyllococcus Ashmead, 1879. Canad. Ent. 11:160. Dactylopius destructor Comstock, 1880-1881. Rept. U.S. Dept. Agr., p. 342. Dactylopius brevispinus Targioni, 1881. Min. d'Agr., Indus, e Com., Ann. di Agr. 34:137. Pseudococcus calceolaria var. minor Maskell, 1897. Trans. New Zeal. Inst. 29:322. Pseudococcus citri (Risso), Fernald, 1903. Mass. Agr. Expt. Sta. Spec. Bull. 88, p. 99. Pseudococcus citri var. phenacocciformis Brain, 1915. Trans. Roy. Soc. So. Africa 5 ( 2 ) : 116. Pseudococcus citri (Risso), Ferris, 1918. Stanford Univ. Pubs., Univ. Ser. pp. 39-40, illus. Planococcus citri (Risso), Ferris, 1950. Atlas of scale insects of North America, Ser. V: 165-167, illus. Planococcus citri (Risso), Ezzat and McConnell, 1956. Md. Agr. Expt. Sta. Bull. A-84 (Tech.), pp. 65-67, illus. Planococcus citri (Risso), Williams, 1962. Bull. Brit. Mus. (Nat. Hist.) Ent. 12(1) :39. Type Locality and Host. Southern France, on Citrus sp. (Rutaceae). North American Distribution. Occurring very commonly in greenhouses in all of North America. In the United States it thrives out-of-doors in the southern states and in southern California. Additional Hosts. The host list of this species is endless, and the species may be expected to occur on almost any flowering plant. The following list is a sample to show diversity of hosts infested by this mealybug: Ananas sativus (= A. cumunis) (Bromeliaceae), Annona squamosa (Annonaceae), Asparagus sp. (Liliaceae), Bougainvillea sp. (Nyctaginaceae), Cactus sp. (Cactaceae), Citrus spp. (Rutaceae), Cocos nucifera (Palmaceae), Coleus sp. (Labiatae), Codiaeum sp. (Euphorbiaceae), Cucumis

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OF

SPECIES

Fig. 110. Planococcus citri (Risso), collected on lemon, Citrus limon (Rutaceae), at Azusa, Los Angeles County, California. (Illustration prepared from specimens collected on Coleus sp. [Labiatae], at Ontario, San Bernardino County, California.)

281

282

TAXONOMY

OF

SPECIES

melo (Cucurbitaceae), Cucurbita sp. (Cucurbitaceae), Cyperus sp. (Cyperaceae), Dioscorea sp. (Dioscoreaceae), Euphorbia sp. (Euphorbiaceae), Ficus sp. (Moraceae), Fragaria sp. (Rosaceae), Gardenia sp. (Rubiaceae), Impatiens sp. (Balsaminaceae), Ipomoea batatas (Convolvulaceae), Mangifera indica (Anacardiaceae), Musa sp. (Musaceae), Nicotiana sp. (Solanaceae), Oryza latifolia (Gramineae), Persea americana (Lauraceae), Phaseolus limenis (Leguminosae), Phoenix dactylifera (Palmaceae), Psidium guava (Myrtaceae), Punica granatum (Punicaceae), Pyrus communis (Rosaceae), Pyrus malus or Malus sylvestris (Rosaceae), Rosa sp. (Rosaceae), Solanum melongena (Solanaceae), and Theobroma cacao (Sterculiaceae). External Features and Habitat. The citrus mealybug is quite thickly covered with a powdery, waxy secretion, and the lateral wax filaments are very short, with the caudal pair only slightly longer than the others. A fluffy white ovisac is formed at maturity (see col. pis. I and XIV). The species ordinarily occurs on the foliage or stems of its host. Recognition Characters. Adult female, mounted, 2.00 to 3.50 mm long, 1.10 to 2.00 mm wide; body form oval. Dorsum normally with 18 pairs of cerarii. Anal lobe cerarius with 2 rather slender, conical setae, 3 or 4 slender auxiliary setae, small group (about 20) associated trilocular pores, and usually light sclerotization. Remaining cerarii likewise with but 2 conical setae, these slightly smaller than those on anal lobes, no auxiliary setae, each with very small cluster trilocular pores, no sclerotization. Dorsum with trilocular pores abundant, evenly distributed. Minute circular pores scattered. Tubular ducts large, of a modified oral-collar type (appearing as oral-rim tubulars in well-stained examples), either lacking or very few, a single duct may be present near any cerarius on eighth to second abdominal segments, more frequent on eighth to sixth segments, with 1-4 ducts scattered on thorax and head. Dorsal body setae all small, slender, occasionally very slightly lanceolate, never much swollen toward base. Anal ring apical, with no unusual features; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores numerous from area posterior to vulva forward to posterior margin of fourth abdominal segment; occurring in very small numbers in thoracic sternal areas. Trilocular pores rather numerous, though quite evenly distributed over entire venter. Minute circular pores scattered. Small oral-collar tubular ducts present, especially in clusters near lateral border of abdominal segments; scattered along margin of thorax. Body setae slender, some slightly longer than those on dorsum. Ventral surface with sharply defined, narrow, sclerotized bar extending in from base of anal lobe seta, and bearing near its center a smaller seta. Circulus fairly large, transversely oval, capable of folding along intersegmental line. Legs comparatively large;

translucent pores present on coxa and tibia; claw without denticle or tooth on plantar surface. Antennae 8-segmented, slender. Notes. This is the so-called citrus mealybug of economic entomologists. In the United States, it occurs out-of-doors in the southern states and in southern California. In California, it is of importance on Citrus. In greenhouses, it occurs especially on Coleus. This species is closely related to Planococcus kraunhiae (Kuwana), the only other known species of this group in California, but differs in having on the dorsum usually only a single oral-collar tubular duct (these often with slight oral-rim indicated) near any cerarius on abdomen, whereas in the species last mentioned there are groups of these structures near the abdominal cerarii. P. citri also has slender dorsal body setae while those of P. kraunhiae are obviously swollen at base. California Records (Map 101) Alameda Co.: Berkeley (U. C. Campus), IV20-22, on Citrus sp. (collector ?, CAS); Berkeley, XII-24-58, on Dieffenbachia sp. (D. Baer, UCD); San Leandro, XI-10-48, on Stephanotis sp. (G. B. Laing, CDA). Fresno Co.: Fresno, VII-20-42, on Ceratonia siliqua (J. W. Dixon, CDA); VI-22-60 and VI-23-60, on ? (H. V. Dunnegan and J. Perry, CDA). Humboldt Co.: Fortuna, X-12-60, on ? (K. E. Miller, CDA). Imperial Co.: Brawley, VIII-15-59, on Hibiscus sp. (E. I. Schlinger, UCD). Kern Co.: Bakersfield, I X - 3 0 ^ 1 , on Wisteria sp. (J. W. Dixon, CDA). Los Angeles Co.: Alhambra, X-28-12, on Magnolia sp. (R. S. Woglum, USNM); 1918, on Citrus sinensis (R. S. Woglum, UCD); Arcadia, V-15-63, on Sciaphyllum sp. (R. Wightman, UCD); Azusa, XII-1921, on Citrus limón (G. F. Ferris, UCD);Los Angeles, IV-4-32, on Franciscia sp. (J. H. Caldwell, CDA); Pacific Palisades, V-13-38-, on Pinus sp. (dwarf) (L. E. Myers, CDA); Pasadena, V-25-16, on Citrus sinensis (R. S. Woglum, USNM); San Dimas, 1-28-27, on "carob" (L. E. Myers, L E M ) ; Whittier, III-6-34, on Citrus sinensis (H. Compere, UCD). Marin Co.: Mill Valley, IV-8-42, on Stephanotis sp. (M. R. Bell, CDA). Monterey Co.: Salinas, X-18-44, on Amaryllis sp. (M. R. Bell, CDA). Orange Co.: Anaheim, X-25-38, on Citrus sp. (R. J. Bumgardner, CDA); Brea, IX-27-40, on Citrus limón (C. E. Norland, CDA). Riverside Co.: Gilman Hot Springs, XI-13-40, on Coleus sp. and Euphorbia pulcherrima (C. R. Tower, CDA); XI-20-42, on Amaryllis sp. (hybrid) (C. E. Ehmann, CDA); Locality ?, 1-8-43, on Citrus sp. (orange) (C. Z. Yonge, CDA). Sacramento Co.: Carmichael, 1-25-59, on Saintpaulia sp. (M. L. Jones, CDA); IX-19-62, on Pinus palustris (J. C. Wilson, CDA); Sacramento, X-13-34, on Coleus sp. (B. Callins, CDA); III-3-33, on

COLOR PLATE XIV. The citrus mealybug, Planococcus citri (Risso), on col»us, Coleus sp. (Cruciferae), Davis (University of California greenhouse), Yolo County, California.

Choisya sp. (E. T. Gammon, CDA) ; VIII-18-63, on Acorus sp., Gloxinia sp., and Hoya sp. (H. L. McKenzie, UCD). San Bernardino Co.: Ontario, VI-19-30, on Coleus sp. (G. A. Pohl, UCD). San Diego Co.: Chula Vista, X-18-11, on Citrus limon (P. H. Timberlake, USNM); 1918, on Citrus limon (R. S. Woglum, UCD); IV-12-32, on Citrus sp. (orange) (S. E. Flanders, UCD) ; Coronado, VIII-30-12, on Cupressus sp. (P. H. Timberlake, USNM) ; El Cajon, XI-28-36, on Citrus sp. (orange) (J. O. Brodner, CD A ) ; Escondido, 1918, on Citrus limon (R. S. Woglum, UCD); San Diego, X-3-34, on Coleus sp. (F. Aller, CDA); III-1920, on Solanum sp. (roots) (G. R. Groton, UCD); III-4-32, on Solanum sp. (potato) (L. E. Myers, CD A). San Francisco Co.: San Francisco, VIII-30-06, on Nerium oleander (O. E. Bremner, UCD). San Joaquin Co.: Manteca, VIII-25-59, on Ipomoea batatas (stored) (J. L. Joos, UCD). San Mateo Co.: Colma, VII-29-29, on Gardenia sp. (F. W. McLelland, CDA). Santa Barbara Co.: Santa Barbara, X-7-11, on Citrus limon (collector ?, UCD); VII-29-43, on Strelitzia regina (C. E. Ehmann and R. P. Allen, CDA). Santa Clara Co.: Stanford University, XI-27-16, on Begonia sp. (in greenhouse) (G. F. Ferris, UCD). Solano Co.: Vacaville, VI-4-62, on Strelitzia sp. (I. W. Price, CDA). Stanislaus Co.: Modesto, X-22-31, on Coleus sp. (E. T. Hamlin, CDA); Turlock, VII-11-62, on Ficus pandoratus (G. Wilhite, CDA). Tehama Co.: Locality ?, 1918, on Coleus sp. (H. S. Smith, UCD). Ventura Co.: Fillmore, 1918, on Citrus limon (R. S. Woglum, UCD); Santa Paula, XII-1921, on Citrus sinensis (A. H. Call, UCD) ; X-5-09, on Citrus sp. (R. S. Woglum, USNM); XI-8-08, on Cucurbita pepo (P. E. Smith, USNM); date ?, on Solanum douglasii (roots) (collector ?, CAS); no specific locality, IX-1962, on Citrus limon (J. C. Ortega, UCD). Yolo Co.: Davis, VII-8-58, on Apium graveolens (A. A. Grigarick, UCD) ; XI-1958, on Beta vulgaris (A. A. Grigarick, UCD); 11-23-60, on Coleus sp. (H. L. McKenzie, UCD); X-19-62, on Medicago sativa (in greenhouse) (W. M. Gilbert, UCD); V I 30-58, on Philodendron sp. (W. G. Iltis, UCD); I V 7-59, on Populus trichocarpa (A. A. Grigarick, UCD); V-13-59, on Vitis sp. (R. O. Schuster, UCD); Davis, U.C. Greenhouse, XI-5-64, on Beta vulgaris (foliage) (D. R. Miller, UCD). Yuba Co. : Marysville, date ?, on Olea europaea ? (collector ?, UCD).

Map 101. Distribution in California of Planococcus citri (Risso).

Planococcus kraunhiae (Kuwana) Japanese Mealybug (Figure 111) Dactylopius kraunhiae Kuwana, 1902. Proc. Cal. Acad. Sci. (Ser. 3, Zoology) (2):55-56, illus. Pseudococcus kraunhiae (Kuwana), Fernald, 1903. Mass. Agr. Expt. Sta., Spec. Bull. 88, p. 104. Pseudococcus kraunhiae (Kuwana), Ferris, 1918. Stanford Univ. Pubs., Univ. Ser. pp 45-46, illus. Planococcus kraunhiae (Kuwana), Ferris, 1950. Atlas of scale insects of North America, Ser. V: 168-169, illus. Planococcus kraunhiae (Kuwana), Ezzat and McConnell, 1956. Md. Agr. Expt. Sta. Bull. A-84 (Tech.), pp. 87-89, illus. Type Locality and Host. Yokahama (nursery), Japan, on Japanese wisteria, Wisteria floribunda (= Kraunhia floribunda) (Leguminosae). North American Distribution. Known only from California. Additional Hosts. Citrus nobilis var. unshi (Rutaceae), C. paradisi (Rutaceae), Diospyros sp. (Ebenaceae), Ficus carica (Moraceae), Ilex sp. (Aquifoliaceae), Olea chrysophylla (Oleaceae), and Plantanus orientalis (Platanaceae). External Features and Habitat. According to Ferris (1950), this mealybug is "apparently very similar to citri, but differing in having red body contents." The species presumably occurs on the foliage and stems of its host. Recognition Characters. Adult female, mounted, 2.00 to 2.70 mm long, 1.00 to 1.70 mm wide; body form oval. Dorsum normally with 18 pairs of cerarii. All cerarii, except perhaps head pairs, with 2 rather slen-

284

TAXONOMY

OF

SPECIES

Fig. 111. Planococcus kraunhiae (Kuwana), collected on Citrus sp. (Rutaceae), at Ojai, Ventura County, California. (Illustration prepared from specimens collected on Wisteria sp. [Leguminosae], at Ojai, Ventura County, California.)

TAXONOMY

der conical setae, these much the same size throughout, tending to be slightly flagellate at apex, never with slender auxiliary setae, except on anal lobes, accompanied in each case by very small group trilocular pores, no sclerotization. Dorsum with rather numerous trilocular pores quite uniformly beset over surface. Minute circular pores lightly scattered. Tubular ducts of oral-collar type present, small, orifice slightly larger than trilocular pore, difficult to find except in well-stained specimens, 1-4 near each abdominal cerarius of eighth to second segments, few situated in more or less transverse row across most abdominal segments. Dorsal body setae variable but tending to be noticeably large, swollen toward base, apex more or less flagellate. Anal ring apical, with no unusual characters; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores abundant from ninth abdominal segment forward to fifth and in lateral areas, occasional pore situated in thoracic area. Trilocular pores quite uniformly beset on venter. Very small oral-collar tubular ducts numerous in midregion of abdomen, in clusters in lateral areas of abdominal segments, extending around to dorsal surface, scattered on thorax and head. Body setae slender, some longer than those on dorsum though differently shaped. Ventral surface with definite, narrow, sclerotized bar extending in from base of anal lobe seta, and bearing near its center a smaller seta. Circulus quite large, transversely oval, divided by segmental line between which it lies. Legs comparatively large, translucent pores present on coxa and distal end of tibia; claw without denticle on plantar surface. Antennae 8-segmented. Notes. This species is similar to Planococcus citri (Risso), but differs from it by the presence of groups of dorsal oral-collar tubular ducts situated near abdominal cerarii, and by the character of the dorsal body setae which are obviously swollen at base. P.

SPECIES

285

citri, on the other hand, has only a single dorsal oralcollar duct situated near any cerarius on abdomen, and the dorsal body setae are basically slender (not swollen) throughout their entire length. California Records (Map 102) Santa Barbara County: No specific locality, 1926, on Citrus sp. (A. Call, UCD). Ventura Co.: Nordhoff, IX-10-15, on Wistaria sp. and Diospyros sp. (A. H. Brook, CDA); Ojai, 1918, on Citrus sp. (H. S. Smith, UCD); IV-23-43, on Wistaria sp. (collector ?, UCD); VII-27-48, on Wistaria sp. (collector ?, CDA).

Map 102. Distribution in California of Planococcus kraunhiae (Kuwana). Marin-* SmI Frttiieitee SonMotto

PSEUDANTONINA Pseudantonina Green, 1922. The Coccidae of Ceylon (London: Dulau), part V:363. Pseudantonina Green, Ferris, 1953. Atlas of the scale insects of North America, Vol. VI: 411. Genotype. Pseudantonina bambusae Green. Recognition Characters. Pseudococcidae with body form from rather narrowly oval to broadly oval or rotund. Dorsal ostioles recognizable only in posterior pair in some species, both pairs present in other species. Cerarii lacking entirely or represented only by anal lobe pair containing 2 very small conical setae. Multilocular disk pores and trilocular pores commonly present. Tubular ducts of various forms present, none, however, of typical pseudococcid form.

OF

GREEN

Anal ring large, broad, strongly sclerotized, equipped with oval pores, at times somewhat invaginated into body. Circulus lacking. Legs greatly reduced in size, at times somewhat distorted, retaining usual parts; posterior coxa either enormously enlarged and flattened into a plate or area beset with small pores, or coxa normal in size and lying in an enlarged area beset with small pores; claw without denticle or tooth on plantar surface. Notes. Distinctive features of this genus is the area of pores borne on an enlarged, flattened coxal plate or on area surrounding this plate, combined with the reduced legs and a 5- or 6-segmented antenna.

286

TAXONOMY

OF

SPECIES

Three species of Pseudantonina are present in North America, 1 of which occurs in California. All of these species infest grasses.

KEY Species of Pseudantonina in North America: Adult Females (Modified from Ferris, 1953)

1.

2(1).

Multilocular disk pores scattered over entire dorsum giganticoxa Hoke Multilocular disk pores confined to venter except for a few that may occur on dorsum along lateral margins of terminal abdominal segments 2 Multilocular disk pores on venter present as a submarginal band from posterior apex of abdomen forward to metathorax . .. arundinariae McConnell Multilocular disk pores on venter situated mainly in midregion of seventh to tenth abdominal segments texana Ferris

Pseudantonina arundinariae McConnell Arundinaria Mealybug (Figure 112) Pseudantonina arundinariae McConnell, 1941. Proc. Wash. Ent. Soc. 43(5):93-95, illus. Pseudantonina arundinariae McConnell, Ferris, 1953. Atlas of scale insects of North America, Vol. VI:412-413, illus. Type Locality and Host. Anderson, Anderson County, South Carolina, on Arundinaria sp. (Gramineae). North American Distribution. California and South Carolina. Additional Hosts. "Bamboo" (Gramineae). External Features and Habitat. McConnell (1941) gives the following account of this mealybug in his original description: "Adult female, in life, occurring between leaf sheaths of host, flattened on account of position, derm pink to reddish (dead specimens dark brown to black), dorsum nearly naked, powdery wax along margins, especially posterior part of abdomen; no definite ovisac formed, eggs pushed out behind and mixed with powdery wax." Recognition Characters. Adult female, mounted, 2.50 to 5.20 mm long, 1.70 to 2.50 mm wide; body form slightly irregular in shape, narrowly oval, sides subparallel, tapering somewhat toward well-rounded ends of body, derm slightly sclerotized at margins. Apparently only posterior pair of ostioles present. Dorsum with anal lobe cerarius alone present, these each with 2 very small, conical setae. Dorsal multilocular disk pores present in submarginal patches from ninth abdominal segment to penul-

timate and antepenultimate areas. Trilocular pores evenly beset over entire dorsal surface. Tubular ducts short, lightly scattered on dorsum, absent on head. Body setae small and slender. Anal ring of normal size, strongly sclerotic, bearing 2 rows of oval pores; each of its 6 setae perhaps one and a half times as long as diameter of ring itself. Venter with multilocular disk pores abundant from posterior apex of abdomen to segment 7, and in lateral zone extending forward onto head. Trilocular pores sparingly beset on ventral surface. Tubular ducts short, especially noticeable on last 4 or 5 abdominal segments, and intermixed with submarginal band of multilocular disk pores. Body setae small and slender, some longer than those on dorsum. Circulus absent. Spiracles moderately sclerotized, with several trilocular pores loosely grouped lateral to orifice. Legs very small and slender, probably nonfunctional; coxae lying in an enlarged area beset with small pores. Antennae small, 6-segmented. Notes. This remarkable pseudococcid is related to Pseudantonina texana Ferris, a Texas species, but differs from it mainly in having coxae of hind legs normally shaped and developed, whereas, in the species last mentioned, the posterior coxae are greatly expanded and flattened into a large plate bearing many small pores. California Records (Map 103) Los Angeles Co.: Baldwin Park, II-5-59, on "bamboo" (dwarf) (L. Johnson, CD A ) : III-3-59, on "bamboo" (dwarf) (L. Johnson, CDA).

TAXONOMY

OF

SPECIES

Fig. 112. Pseudantonina arundinariae McConnell, collected on "bamboo" (dwarf) (Gramineae), at Baldwin Park, Los Angeles County, California. (Illustration prepared from type material.)

287

288

TAXONOMY

OF

SPECIES

PSEUDOCOCCUS Pseudococcus Westwood, 1840. An introduction to the modern classification of insects, founded on the natural habits and corresponding organization of different families, Vol. II, pp. 443-450 (448). Synopsis of the British Insects (at end of Vol. II), 158 pp. (118). Pseudococcus Westwood, Ferris, 1950. Atlas of the scale insects of North America, Ser. V: 170-171. Genotype. Pseudococcus longispinus (Targioni Tozzetti) (pending permission of the International Commission on Zoological Nomenclature). Recognition Characters. Pseudococcidae, for the most part, with body form at maturity usually broadly oval to subcircular. Two pairs of dorsal ostioles. Normally not more than 17 pairs cerarii, at times with less, these usually with but 2 conical setae, at times with more; normally at least in part with slender auxiliary setae; usually with numerous trilocular pores either clustered or scattered. Dorsum often with discoidal and minute circular pores present. Oral-rim tubular ducts normally present on either dorsal or ventral surface. Oral-collar type tubular ducts may occur on both body surfaces, though most commonly on venter. Anal ring usually large and cellular, bearing 6 setae each approximately twice as long as diameter of ring. Venter always with at least a few multilocular disk pores. Quinquelocular pores absent. Circulus present or absent, when present often extending across fold between fourth and fifth abdominal segment. Legs always well developed; translucent pores rather common on coxae and tibiae; claw without denticle or tooth on plantar surface. Antennae normally 8-segmented. Notes. This genus has close affinities with Dysmicoccus Ferris. However, the consistent pattern of the dorsal oral-rim tubular ducts in Pseudococcus (one species has them restricted to venter), combined with other characters enumerated above, seems actually to distinguish it quite satisfactorily from Dysmicoccus. Two new California species of Pseudococcus are described in this book. To the genus there are now assigned 20 species, 14 of which occur in California.

KEY Species of Pseudococcus in North America: Adult Females

1.

2(1).

With at least a few translucent pores present on trochanter of hind leg . . . sorghiellus (Forbes) Translucent pores absent on trochanter of hind leg 2 Circulus small, transversely oval, usually on fourth abdominal segment,

WESTWOOD never divided by intersegmental line, even if situated across it microcirculus McKenzie Circulus normally quite large and, even if small, always divided by line between fourth and fifth abdominal segments 3 3(2). With at least 1 oral-rim tubular duct on dorsum 4 Without oral-rim tubular ducts on dorsum diversus McKenzie 4 ( 3 ) . With not more than 1 oral-rim tubular duct near any cerarius 5 With 2 or 3, usually 3, oral-rim tubular ducts near almost every cerarius . . . longispinus (Targioni Tozzetti) 5(4). With 1 or more small, oval, discoidal pores along margin of eye 6 Without discoidal pores along margin of eye 8 6(5). Femur of hind leg with from 35-81 translucent pores somewhat scattered over segment obscurus Essig Femur of hind leg normally without translucent pores 7 7 ( 6 ) . Dorsal submarginal oral-collar tubular ducts present on last 6 or 7 abdominal segments; numerous ventral oral-collar tubular ducts of 1 size on abdominal segments; a field-inhabiting pseudococcid pertusus McKenzie Dorsal oral-collar tubular ducts absent (infrequently 1 or 2 present); comparatively few ventral oral-collar tubular ducts of 2 sizes on abdominal segments; a nursery species infesting orchids importatus McKenzie 8(5). Certain dorsal body setae on abdomen enlarged, conical, strongly resembling cerarian conical setae, some as large as or larger than those in the cerarii megasetosus McKenzie Dorsal body setae on abdomen slender, not conical-shaped 9 9(8). With not more than 7 oral-rim tubular ducts on dorsum of abdomen 10 With at least 10 oral-rim tubular ducts on dorsum of abdomen, usually more 12 10(9). With 17 pairs of cerarii 11 With fewer than 16 pairs of cerarii . . . aberrans McKenzie 11(10). Discoidal-type pores scattered on dorsal and ventral body surfaces; venter of

TAXONOMY

12(9).

13(12).

14(13).

15(13). 16(15).

17(15).

18(17).

19(18).

sixth abdominal segment without multilocular disk pores, or sometimes with 1 such pore; at least 1 or 2 oralrim tubular ducts on dorsum of abdomen; body form subcircular, as mounted; found on Prunus sp prunicolus McKenzie Discoidal-type completely absent; venter of sixth abdominal segment with 3 to 5 multilocular disk pores along posterior margin; no oral-rim tubular ducts on dorsum of abdomen; body form broadly oval, as mounted; found on Agave sp agavis MacGregor Dorsum with 1 oral-rim tubular duct just posterior to each frontal cerarius 13 Never with an oral-rim tubular duct in this position fragilis Brain Multilocular disk pores on venter of abdomen on only the last 3 or 4 segments 14 Multilocular disk pores on venter of last 15 5 or more abdominal segments With 40 or more multilocular disk pores on venter, on anterior and posterior margins of eighth abdominal segments; no ventral oral-rim tubular ducts near anterior spiracles dispar McKenzie With 25 or fewer multilocular disk pores on venter, on anterior and posterior margins of eighth abdominal segment; a few ventral oral-rim tubular ducts near anterior spiracles sparsus McKenzie With 16 or 17 pairs of cerarii 17 With not more than 15 pairs of cerarii. . 16 Multilocular disk pores on venter of all abdominal segments macswaini McKenzie Multilocular disk pores on venter only from apex of abdomen to posterior margin of fifth segment pseudobscurus McKenzie With translucent pores on hind coxae . . cornstocki (Kuwana) Without translucent pores on hind coxae 18 Dorsal body setae strikingly long, some as long as the anterior posterior measurement of the segment that bears them, or longer . . . . longisetosus Ferris All dorsal body setae short, none as much as half this measurement . . . . 19 With more than 20 oral-rim tubular ducts on dorsum of thorax and head maritimus (Ehrhorn) With fewer than 10 oral-rim tubular ducts on dorsum of thorax and head kingii (Cockerell)

OF

SPECIES

289

Pseudococcus aberrans McKenzie Aberrant Mealybug (Figure 113) Pseudococcus aberrans McKenzie, 1962. Hilgardia 32(14):656-658, illus. (nec. Trionymus aberrans Goux, 1938). Type Locality and Host. Compton, Los Angeles County, California, on juniper, Juniperus torulosa (Cupressaceae). North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. No available information on external features of this mealybug. The mealybug presumably feeds on the foliage and stems of its host. Recognition Characters. Adult female, mounted, 1.80 to 2.50 mm long, 1.15 to 2.00 mm wide; body form broadly oval. Dorsum with cerarii reduced in number for members of this group, 12 to 13 pairs observed, tendency being for cerarii to disappear along thoracic area. Anal lobe cerarius with 2 quite large and stout, conical setae, 2 or 3 slender auxiliary setae, concentrated cluster trilocular pores, very slight suggestion of sclerotization. Penultimate cerarius with 2 smaller, conical setae, 1 or 2 slender auxiliary setae, few trilocular pores, without sclerotization. Anterior to this, cerarii with conical setae progressively smaller, variable in number, ranging from 2-4 (greater number especially noticeable anteriorly), auxiliary setae present or absent, fewer accompanying trilocular pores, without sclerotization. Trilocular pores evenly distributed on dorsum. Minute circular pores scattered on body. Moderately large oral-rim tubular ducts sparsely distributed over dorsum, only 2 or 3 of these structures observed on abdomen, fewer on thorax and head. Dorsal body setae slender, proportionately longer than usual, practically as long as those in same area on venter. Anal ring of normal form and size for genus; each of its 6 setae at least twice as long as greatest diameter of ring. Venter with multilocular disk pores fairly numerous, occurring on last four abdominal segments, absent on thorax and head. Trilocular pores fairly evenly distributed on venter. Minute circular pores scattered over entire ventral surface. Oral-collar tubular ducts essentially of 1 size, appearing quite abundantly along submargins of abdomen and on posterior margin of most abdominal segments. Oralrim tubular ducts fairly numerous, situated laterally on third abdominal segment, submarginally between middle and hind legs, few near anterior spiracles, absent on head. Body setae slender, slightly longer than those of dorsum. Circulus quite large, normally appearing broadly oval, somewhat compressed, capable of folding along intersegmental line across which it lies. Legs relatively large, well developed, femora and tibiae moderately

290

TAXONOMY

OF

SPECIES

Fig. 113. Pseudococcus aberrans McKenzie, collected on juniper, Juniperus torulosa (Cupressaceae), at Compton, Los Angeles County, California.

TAXONOMY

swollen; entire length of hind tibiae quite numerous translucent pores, 41 to 50 of these structures observed; claw without denticle or tooth on plantar surface. Antennae 8-segmented, slender. Notes. Borchsenius (1949) inadvertently transferred Trionymus aberrans Goux, described in 1938, to the genus Pseudococcus. According to the International Rules of Nomenclature, this would place my species as a secondary homonym were it not for the fact that Goux' aberrans is unquestionably a member of Trionymus. It is here returned to that group. Pseudococcus aberrans McKenzie appears related to P. diversus McKenzie (a mealybug infesting cypress in California), but may be distinguished from it mainly by the presence of at least a few dorsal oralrim tubular ducts situated along submargin of abdomen, on thorax and head; and oral-collar tubular ducts especially abundant along submargin of abdominal segments 8-7. P. diversus, on the other hand, lacks dorsal oral-rim tubulars anywhere on dorsal surface; and has only a few oral-collar ducts along submargin of abdominal segments 8-7. California Records (Map 104) Los Angeles Co.: Compton, VI-8-60 and XI-2-61, on Juniperus torulosa (J. H. Caldwell, CDA). Riverside Co.: Banning (nursery), X-22-65, on Juniperus sp. (N. W. Getz, CDA). San Diego Co.: San Diego, IV-1-57, on Cupressus sp. (R. Brown, CDA).

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Puto bryanthi Ferris Bryanthus Mealybug (Figure 133) Puto bryanthi Ferris, 1950. Atlas of scale insects of North America, Ser. V: 199-200, illus. Type Locality and Host. Above Mineral King (9,500-10,000' elev.). Tulare County, California, on red heather, Phyllodoce (=Bryanthus) breweri. (Ericaceae). North American Distribution. California, South Dakota, and Wyoming. Additional Hosts. None. External Features and Habitat. According to Ferris (1950), "The host is a mat plant, and the insects occur on the stems well down in the mat." No other information is available at this time. Recognition Characters. Topotype adult females, certainly mature but probably not fully expanded, 2.20 to 3.10 mm long, 1.40 to 2.00 mm wide, adult females from South Dakota and Wyoming, probably fully developed and extended, 3.00 to 4.10 mm long, 1.99 to 3.00 mm wide; body form broadly oval at maturity. Dorsum basically with 18 pairs of cerarii, this arrangement, however, interrupted by division of in-

OF

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dividual cerarii or interpolation of others; all cerarian plates quite small, sclerotized, each plate bearing from 2 to as many as 16, or perhaps even more, enlarged setae, these apically pointed, slightly lanceolate, tubular ducts completely absent, trilocular pores borne on plates usually more numerous than setae. Trilocular pores rather evenly distributed on dorsum. Dorsal body setae as large as or only slighdy smaller than the cerarian setae, relatively few, arranged in small clusters or short, interrupted rows, basal scleroses absent. Anal ring apical, typical for genus; each of its 6 setae nearly twice as long as greatest diameter of ring. Venter with multilocular disk pores of 2 very slightly different sizes, with more or less sclerotized outer ring, situated predominantly in. midregion of body from posterior apical abdominal segment forward to head, those anterior to posterior margin of seventh segment slightly smaller. Trilocular pores relatively sparse over venter, certain "clear" areas especially noticeable in sternal region. Oral-collar tubular ducts situated across and in a submarginal band on most abdominal segments; several more slender and elongated ones noticeable in head area anterior to mouthparts. Body setae long and slender, for most part exceeding in length those on dorsum. Circulus quite small, broadly oval. Legs fairly long; claw with prominent denticle on plantar surface. Antennae normally 9-segmented, slender. Notes. This species is quite similar to Puto ambiguus (Fullaway) and to P. barberi (Cockerell), agreeing with both in the absence of tubular ducts on the cerarian sclerotized plates. It may be separated from these 2 species by the shape of its circulus, this being extremely small for members of this genus and short-oval along its transverse diameter, in contrast with a transversely elongated circulus of the other 2. In addition, Ferris (1950) maintains that on ecological grounds alone it would seem highly probable that P. bryanthi is distinct from the other 2 species since it occurs at very high elevations, whereas the others are found near sea level. California Records (Map 124) Mono Co.:Rock Creek (10,500' elev.), near Half Moon Pass, IX-22-61, on moss (E. I. Schlinger, UCD). Tulare Co., Above Mineral King (9,500-10,000' alt.), VIII-29-46, on Phyllodoce (=Bryanthus) breweri (G. F. Ferris, UCD).

Puto californicus McKenzie, new species California Puto Mealybug (Figure 134; Color Plate IV) Type Locality and Host. Type and paratype adult females on deerweed, Lotus scoparius (Leguminosae), 5 miles north of San Bernardino, San Bernardino County, California, collected June 11, 1963, and July 23, 1963, by D. R. Miller. Additional paratype adult females are available on perennial eriastrum, Eriastrum (=Hugelia) densifolium (Polemoniaceae),

340

TAXONOMY

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Fig. 133. Puto bryanthi Ferris, collected on red heather, Phyllodoce (= Bryanthus) breweri (Ericaceae), at Mineral King (9,500-10,000 feet elevation), Tulare County, California.

TAXONOMY

OF

SPECIES

Fig. 134. Puto californicus McKenzie, new species, collected on deerweed, Lotus scoparius (Leguminosae), north of San Bernardino, San Bernardino County, California.

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Sah BamarJina &nnùda AmPftfo [mprial 1 mile south Cajon, San Bernardino County, California, collected June 24, 1964, by D. R. Miller; and on various "wild plants," Uplands, San Bernadino County, California, collected July 1, 1914, by C. P. Clausen. Type Material. Holotype adult female (single specimen on slide) and female paratypes in museum, University of California, Davis. Additional paratypes in United States National Collection of Coccoidea (U.S. National Museum), Washington, D. C., and in collection of California State Department of Agriculture, Sacramento. North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. This mealybug is heavily covered with a white, fluffy, wax secretion. Its body form is subcircular; body color variable from pink to green (green being most common); body contents when crushed are grayish; and legs dark brown. There are waxy filaments around the body, caudal pair twice as long as all the rest (see col. pi. IV). No ovisac is produced. The male is winged. This species occurs on the crown and roots of its host. Ants were found associated with the mealybug nymphs on 1 occasion. Recognition Characters. Adult females, mounted, 4.00 to 6.00 mm long, 2.60 to 3.60 mm wide; body from broadly oval. Dorsum with basically 18 pairs of cerarii, this arrangement, however, interrupted by division of individual cerarii or interpolation of others; each cerarius formed upon a sclerotized plate bearing from 2 or 3 to as many as 25, or perhaps even more, enlarged setae, these apically pointed and slightly lanceolate, trilocular pores present in numbers about equal to setae. Trilocular pores fairly evenly distributed on dor-

sum. Modified minute circular or discoidal-type pores scattered over dorsum. Tubular ducts absent on dorsum. Body setae for most part slightly smaller than those of cerarii. Anal ring apical, normal for genus; each of its 6 setae nearly twice as long as greatest diameter of ring. Venter with multilocular disk pores most numerous on last 3 abdominal segments, very few scattered on segment 6 forward to 4 (especially near circulus), near spiracles, and on head. Trilocular pores fairly evenly distributed on venter. Modified minute circular or discoidal-type pores similar to those on dorsum, scattered over venter. Tubular ducts of oral-collar type present in rather small numbers across last 5 abdominal segments, absent elsewhere on venter except for a few more slender, elongated ones situated in cephalic region anterior to mouthparts. Body setae long and slender, greatly exceeding in length those on dorsum. Circulus quite large, broad, transversely oval, with margins appearing somewhat crenulated. Legs unusually large and slender; claw with prominent denticle on plantar surface. Antennae long and slender, normally 9-segmented. Notes. This species is rather closely related to Puto sandini Washburn, but differs in anal lobe cerarius supporting 23-30 (average 25.58) enlarged conical setae, and in being found on representitives of Leguminosae and Polemoniaceae in California. In P. sandini, on the other hand, the anal lobe cerarius possesses only 10-12 (average 11.40) enlarged conical setae, and the species occurs on Picea engelmannii (Pinaceae) in Utah. California Records (Map 125) San Bernardino Co.: 1 mile south Cajon, VI-2464, on Eriastrum densifolium (D. R. Miller, UCD) ; 5 miles north San Bernardino, VI-11-63, on Lotus scoparius (D. R. Miller, UCD); Uplands, VII-1-14, on "wild plants" (C. P. Clausen, UCD). S/$kiifoi TrinihjShasfa j Laxen Tehama } Butte\

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TAXONOMY Puto cupressi (Coleman) California Nutmeg Mealybug (Figure 135) Dactylopius sequoiae Coleman, 1901. Proc. Calif. Acad. Sci (Ser. 3, Zool.), 2(11):412-417, illus. (as to the male). Pseudococcus cupressi Coleman, 1908. Jour. N. Y. Ent. Soc. 16(4): 197-198, illus. Pseudococcus sequoiae Coleman, Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., p. 54 (as to the male). Puto cupressi (Coleman), Ferris, 1950. Atlas of scale insects of North America, Ser. V:203205, illus. Ferris (1950) states: "The description of Dactylopius sequoiae Coleman, from Sequoia sempervirens, at Stanford University, California, was based upon the female of a species of Spilococcus, and the male of the species here under consideration. A type selection was made by Ferris (1918) which restricts this name to the Spilococcus involved." The collection on Pinus ponderosa, Miguel Meadows, Yosemite National Park, California, July 10, 1938, by E. O. Essig and recorded by Ferris (1950) as Puto cupressi is instead P. laticribellum McKenzie. Type Locality and Host. Cypress Point, near Pacific Grove, Monterey County, California, on Monterey cypress, Cupressus macrocarpa (Cupressaceae). North American Distribution. California, Canada, and possibly Utah (last state record based on immature specimens). Additional Hosts. Abies concolor (Pinaceae), Pinus albicaulis (Pinaceae), Picea engelmanni (Pinaceae), Pinus radiata (Pinaceae), Pinus torreyana (Pinaceae), Pseudotsuga taxifolia (Pinaceae), and Sequoia sempervirens (Taxodiaceae). External Features and Habitat. The adult females of this mealybug was described by Ferris (1918) as bare of secretion except for a dusting of fine powder; with short, stout lateral filaments and a somewhat longer caudal pair. He further states that the dorsum is of a light brown color, with marginal and submedian longitudinal rows of dark spots; the ovisac is more or less padlike; and the species is oviparous. Specimens from Abies concolor at Mineral King, California, were enclosed in a fluffy mass of wax threads. The male is winged. According to Ferris (1950), this mealybug occurs' "on either foliage or trunk of the host." The mealybugs on Pinus albicaulis from Lassen Park, California, occurred in a pit beneath the bark, and those from Pinus ponderosa in Yosemite National Park were recorded as occurring under the bark of old trees, near the soil. On Cupressus macrocarpa they may occur about the base of cones. Specimens from Abies concolor, at Mineral King, California, occurred on the needles, and such seems normally to be the case with the species as it occurs on Sequoia. Males seem to outnumber the females, and it is much easier to find male cocoons on Sequoia that it is to find females. Recognition Characters. Adult females, mounted,

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2.60 to 4.50 mm long, 1.60 to 2.50 mm wide; body form broadly oval. Dorsum with basically 18 pairs of very small sclerotized cerarian plates, this arrangement, however, interrupted by division of individual cerarii or interpolation of others; each cerarius with 2 to 17 enlarged setae, or perhaps even more, these apically pointed, slightly lanceolate; within, or just outside margin of most cerarian plates, 1 or more tubular ducts of modified oral-rim type, these rather variable in size, some slightly greater in diameter than diameter of socket at base of cerarian seta, in other cases noticeably larger; trilocular pores borne on plates almost as numerous as setae. Trilocular pores fairly evenly distributed on dorsum. Discoidal type pores lightly scattered on dorsal surface. Tubular ducts of modified oral-rim type sparingly distributed over dorsum. Body setae for most part definitely smaller than those of cerarii. Anal ring apical with usual 6 elongate setae, or with supernumerary, smaller setae, total number of 8 being quite common, infrequently with 10 or more. Venter with quite numerous multilocular disk pores, these all of one size, distributed from posterior apex of abdomen forward to head region. Trilocular pores especially noticeable along submarginal areas of body, apparently absent elsewhere on venter. Discoidal type pores lightly scattered on ventral surface. A few oral-collar tubular ducts present on most abdominal segments, very few scattered on thorax, absent on head. 1 or 2 modified oral-rim tubular ducts present or absent on head. Body setae long and slender, for most part exceeding in length those on dorsum. Circulus transversely elongate oval, margins appearing somewhat crenulated. Legs comparatively short for members of genus; claw with prominent denticle on plantar surface. Antennae normally 8segmented. Notes. The species shares with Puto laticribellum McKenzie possession of enlarged tubular ducts of modified oral-rim type in the cerarii, but differs from the species just mentioned in the absence of conspicuous clusters of ventral submarginal multilocular disk pores on abdomen and near spiracles, a character which readily distinguishes P. laticribellum. California Records (Map 126) Modoc Co.: 6 miles northwest Fort Bidwell, V22-62, on Abies concolor (R. F. Wilkey and T. R. Haig, CDA). Monterey Co.: Monterey (quarantined at Washington, D.C.), V-1918, on ? (C. A. Weigel, USNM); Pacific Grove, XII-1-17, on Pinus radiata (G. F. Ferris, UCD); Pacific Grove (Cypress Point), 1-2504, on Cupressus macrocarpa (G. A. Coleman, UCD); XII-1-17, Cupressus macrocarpa (G. F. Ferris, UCD). San Mateo Co.: Woodside, VII-1-17, on Sequoia sempervirens (G. F. Ferris, UCD). Santa Clara Co.: Los Gatos, IV-3-51, on Aralia sp. (D. F. Zollor, CDA); Stanford University, 1-2117, on Sequoia sempervirens (G. F. Ferris, UCD).

344

TAXONOMY

OF

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Fig. 135. Puto cupressi (Coleman), collected on Monterey cypress, Cupressus macrocarpa (Cupressaceae), at Cypress Point, near Pacific Grove, Monterey County, California.

TAXONOMY

Puto decorosus McKenzie, new species Decorated Mealybug (Figure 136) Type Locality and Host. Type and paratype adult females on Diplacus sp. (Scrophulariaceae), 11 miles southeast Camp Angelus, San Bernardino County, California, collected July 30, 1964, by D. R. Miller. This is the only known collection of the mealybug. Type Material. Holotype adult female (single specimen on slide) and female paratypes in museum, University of California, Davis. Additional paratypes in United States National Collection of Coccoidea (U.S. National Museum), Washington, D. C., and in collection of California States Department of Agriculture, Sacramento. North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. This mealybug is covered with a heavy, white, fluffy, wax secretion. Its body form is subcircular; body color dark green; legs dark brown. There are waxy filaments around body, these quite broad and only moderately long. No ovisac is produced. The male is winged. The mealybug occurs on the roots of its host—on crown only when infestation is heavy. Recognition Characters. Adult females, mounted, 4.30 to 7.10 mm long, 3.00 to 4.75 mm wide; body form broadly oval. Dorsum with basically 18 pairs of cerarii, this arrangement, however, interrupted by division of individual cerarii or interpolation of others; each cerarius formed upon a sclerotized plate bearing from 2 to 38 enlarged setae, or perhaps even more, these apically pointed, somewhat "acorn-shaped," trilocular pores in cerarii about equal in number to setae. Trilocular pores rather numerous and evenly distributed on dorsum. Modified minute circular or dis-

OF SPECIES

345

coidal-type pores scattered in considerable numbers over dorsum. Tubular ducts absent on dorsum. Body setae for most part slightly smaller than those of cerarii. Anal ring apical, normally with 8 enlarged setae, or numbers of these may be increased by smaller supernumerary setae, total of 8 being most common, each of longest setae noticeably knobbed at tip. Venter with numerous multilocular disk pores, these all of approximately one size, distributed from posterior apex of abdomen forward to and including head, noticeable clusters in sternal area of mesothorax, surrounding clypeus of mouthparts, and near spiracles. Trilocular pores fairly evenly distributed on venter. Modified minute circular or discoidal-type pores similar to those on dorsum in shape and distribution. Oral-collar tubular ducts fairly numerous in midregion of posterior apical segment of abdomen forward to segment 5, fewer on segment 4, absent on thorax; a few more slender, elongate tubular ducts situated in cephalic region anterior to mouthparts. Body setae long and slender, exceeding in length those on dorsum, some along lateral margin of abdomen slightly knobbed at tip. Circulus quite large, broad, transversely oval, margins usually slightly crenulated. Legs comparatively short; claw with prominent denticle on plantar surface. Antennae normally 9-segmented. Notes. This species is related to Puto ambiguus (Fullaway), but differs in having anal ring setae, especially largest ones, noticeably knobbed at tip; inhabits mountainous areas; and has a winged male. Puto ambiguus, on the other hand, has anal ring setae lanceolate; occurs in coastal areas; and possesses a wingless male. California Records (Map 127) San Bernardino Co.: 11 miles southeast Camp Angelus, VII-30-64, on Diplacus sp. (D. R. Miller, UCD).

346

TAXONOMY

OF

SPECIES

Fig. 136. Puto decorosus McKenzie, new species, collected on Diplacus sp. (Scrophulariaceae), southeast Camp Angelus, San Bernardino County, California.

TAXONOMY

Puto echinatus McKenzie Fluffy Mealybug (Figure 137 and 138) Puto echinatus McKenzie, 1961. Hilgardia 3 1 ( 2 ) : 35-37, illus. Puto echinatus McKenzie, 1964. Ibid. 35(10) :269. Type Locality and Host. Malibu, Los Angeles County, California, on Batis sp. (Batidaceae). North American Distribution. California and Mexico. Additional Hosts. A triplex semibaccata (Chenopodiaceae). Specimens of this mealybug were collected on the walls of a home at Imperial Beach, San Diego County, California. Subsequent collections on vegetation growing around this home revealed the presence of this pesudococcid on the following hosts: Abronia umbellata (Nyctaginaceae), Chenopodium sp. (Chenopodiaceae), various Compositae, Mesembryanthemum sp. (Arizoaceae), Oenothera cheiranthifolia (Onagraceae), Salsola kali var. tenuifolia (Chenopodiaceae), and Tribulus terrestris (Zygophyllaceae). External Features and Habitat. The adult female of this mealybug is thickly covered with a fluffy, whitish, wax secretion. It possesses short, stout lateral filaments, and- somewhat longer caudal ones. The adult male is wingless. This mealybug infests mainly the crown and roots of its host and is infrequently observed on the lower leaves near the ground. Recognition Characters. Adult female, mounted, 3.40 to 5.00 mm long, 2.00 to 3.60 mm wide; body form broadly oval. Dorsum basically with 18 pairs of cerarii, this arrangement, however, interrupted by division of individual cerarii or by interpolation of others; each cerarius formed upon a sclerotized plate bearing from 8 to 26, or perhaps even more, enlarged conical setae, these apically pointed and slightly lanceolate; 2 to 4 enlarged tubular ducts; trilocular pores usually about as numerous as conical setae, sometimes more so. Trilocular pores rather evenly beset on dorsum. Enlarged tubular ducts of oral-rim type, similar to those in cerarii, variable in number, arranged in groups or short rows on thorax, present or absent on abdomen; if present, very few situated in median and submarginal areas. Dorsal body setae, similar in form to those of cerarii, numerous, arranged in transverse bands on abdominal segments, thorax, and head, many of these setae borne upon well-developed basal scleroses. Anal ring apical, normal for genus; each of its 6 setae nearly twice as long as greatest diameter of ring. Venter with multilocular disk pores, with more or less sclerotized outer ring and all about same size, situated predominantly in midregion of abdomen, from posterior apical to fifth segments, especially numerous on last 3 segments, very few scattered on thorax and head. Trilocular pores rather evenly distributed except for certain "clear" areas in sternal region where they seem to be entirely absent. Oral-

OF

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collar tubular ducts occur in limited numbers on most abdominal segments; a few more slender, elongate ducts situated in cephalic region anterior to mouthparts. Body setae long and slender, for most part exceeding in length those on dorsum. Circulus variable in shape, either broadly oval, or with central portion somewhat oval and prolonged on each side. Legs large, well developed; claw with weakly developed denticle or tooth on plantar surface. Antennae normally 9-segmented. Notes. On the basis of available records, this species seems to be a coastal form commonly associated with Australian saltbush, A triplex semibaccata. It is quite similar to Puto simmondsiae McKenzie, but differs in possessing many dorsal body setae which are borne upon well-developed base scleroses, and by evidence of ventral multilocular disk pores on the fifth and sixth abdominal segments. P. simmondsiae, on the other hand, lacks basal scleroses on any of the dorsal body setae, as well as ventral multilocular disk pores on fifth and sixth abdominal segments. P. echinatus has some affinities with P. yuccae (Coquillett), although the paucity of ventral mutiloculars on thorax and head precludes confusion with the species last mentioned, which has an abundance of these pores in this area. McKenzie (1964) reports that the illustration accompanying the original description of this species was made from a single, poorly-stained, somewhat distorted, holotype adult female. A reexamination of the type slide mount and a critical study of specimens of the species taken in series on A triplex semibaccata from several coastal localities show considerable variations in certain morphological characters, so much so, in fact, that it has been necessary to prepare two separate illustrations to show these differences. Figure 137, is a modification of the illustration accompanying the original description of P. echinatus, altered to include several enlarged, dorsal, oral-rim tubular type ducts on thorax, and a few ventral multilocular disk pores near midcoxae, mouthparts, and base of antennae. Figure 138, made from examples collected along the coast on A triplex semibaccata, shows dorsally a few, enlarged, oral-rim tubular type ducts along median and lateral areas of abdomen; many more of these structures in rows on dorsum of thorax; a circulus that is somewhat produced laterally; and perhaps a few more ventral multilocular disk pores on thorax and head. There is considerable variation in these structures, but aside from this all the specimens at hand hold closely to a common pattern, and the species is here considered quite distinctive. California Records (Map 128) Los Angeles Co.: Malibu, V-17-60, on Batis sp. (A. G. C. Beresford and G. G. Beevor, CD A ) ; Redondo Beach, VI-13-64, on Oenothera cheiranthifolia (D. M. Peterson, UCD). Monterey Co.: Soledad, VIII-15-51, on A triplex semibaccata (W. Lewis and H. T. Osborn, CDA). Orange Co.: Balboa, VIII-1935, on Salsola kali (J. D. Maple, UCD); Newport Beach, IV-14-63 and

348

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Fig. 137. Puto echinatus McKenzie, collected on Batis sp. (Batidaceae), at Malibu, Los Angeles County, California. (Modification of original illustration to include several enlarged, dorsal, oral-rim tubular ducts on thorax; few ventral multilocular disk pores on thorax and head).

TAXONOMY

OF

SPECIES

Fig. 138. Puto echinatus McKenzie, collected on Australian saltbush, A triplex semibaccata (Chenopodiaceae), at Newport Beach (Back Bay), Orange County, California. Note especially the enlarged, dorsal, oral-rim tubular ducts along median and lateral areas of abdomen; and many of these structures in rows on dorsum of thorax.

349

350

TAXONOMY

OF

SPECIES

XI-28-63, on Atriplex semibaccata (D. R. Miller, UCD); Newport Beach (Back Bay), VII-27-63, on Atriplex semibaccata (D. R. Miller, UCD). San Diego Co.: 3 miles south Carlsbad, VII-2763, on Atriplex semibaccata (D. R. Miller, UCD); El Cajon, VII-7-65, on ? (F. Yaruss, CDA); Imperial Beach, IX-6-63, "in house on walls" (R. J. Buckner, CDA); X-10-63, on Abronia umbellata, Chenopodium sp., various Compositae, Mesembryanthemum sp., Salsola kali var. tenuifolia, and Tribulus terrestris (R. J. Buckner and G. Opal, CDA).

Map 128. Distribution in California of Puto echinatus McKenzie.

Puto laticribellum McKenzie Pine Bark Mealybug (Figure 139) Puto laticribellum McKenzie, 1961. Hilgardia 31 (2):37-39, illus. Type Locality and Host. 5 miles north of Old Station, Shasta County, California, on yellow pine, Pinus ponderosa (Pinaceae). North American Distribution. Known only from California. Additional Hosts. Libocedrus decurrens (Cupressaceae) and Pinus jeffreyi (Pinaceae). External Features and Habitat. According to a collector, this mealybug produces a rather weakly developed ovisac. The male is winged. The species infests the cracks of bark on main tree trunk. Recognition Characters. Adult female, mounted, 4.50 to 6.20 mm long, 3.00 to 3.75 mm wide; body form broadly oval.

Dorsum basically with 18 pairs of cerarii, this arrangement, however, interrupted by division of individual cerarii or by interpolation of others; each cerarius formed upon a sclerotized plate bearing 3 to 17, or perhaps more, enlarged conical, apically pointed, slightly lanceolate setae; 0 to 4 tubular ducts of oral-rim type, these apparently absent on thoracic plates; trilocular pores usually somewhat more numerous than conical setae. Trilocular pores distributed rather evenly over dorsum. Tubular ducts, of type represented within cerarian plates, sparingly distributed on abdomen, mostly submarginal. Dorsal body setae, similar in form to those of cerarii, without sclerotized bases, fairly numerous, arranged in transverse bands on abdominal segments, covering thoracic segments and head. Anal ring apical, normally with 8 long, stout, and elongate setae, each nearly twice as long as greatest diameter of ring. Venter with multilocular disk pores in considerable numbers in midregion of abdomen, especially so submarginally on all abdominal segments, except the last, where they appear in dense clusters (whence the specific name was derived), these pores also situated in sternal areas, in dense clusters between spiracles and body margin, and on head. Trilocular pores rather sparingly distributed over venter. Tubular ducts, of 2 slightly different sizes, on most abdominal segments, especially numerous in submarginal areas; few more slender, elongate tubular ducts situated in cephalic region anterior to mouthparts. Body setae long and slender, for most part exceeding in length those on dorsum. Circulus broad, transversely oval, margin somewhat crenulated. Legs proportionately smaller than usual for genus; hind femur and tibia with translucent pores; claw with prominent denticle or tooth on plantar surface. Antennae normally 8-segmented, slender. Notes. The conspicuous clusters of ventral submarginal multilocular disk pores on abdomen and near spiracles immediately set this species apart from all other North American Puto. In the character of cerarian plates enclosing enlarged tubular ducts, this pseudococcid appears related to Puto cupressi (Coleman) and P.yuccae (Coquillett). The collection on Pinus ponderosa, Miguel Meadows, Yosemite National Park, California, July 10, 1938, by E. O. Essig and recorded by Ferris (1950) as Puto cupressi is instead P. laticribellum. Immature specimens of P. laticribellum collected on bark of Pinus ponderosa at Cedar Grove, Fresno County, California, July 13, 1963, by D. R. Miller, were held in a cardboard box (without food) for four months, during which time molting occurred, with the ultimate emergence of fully matured male and female specimens. The adult females were quite small, measuring not more than 2.40 to 3.00 mm long, 1.50 to 1.80 mm wide. California Records (Map 129) Fresno Co.: Cedar Grove, VII-13-63, on Pinus ponderosa (trunk) (D. R. Miller, UCD).

TAXONOMY

OF

Fig. 139. Puto laticribellum McKenzie, collected on yellow pine, Pinus (Pinaceae), north of Old Station, Shasta County, California.

SPECIES

ponderosa

351

352

TAXONOMY

OF

SPECIES

Madera Co.: Bass Lake, V - l l - 3 5 , on Libocedrus decurrens (G. R. Struble, CDA). Mariposa Co.: Yosemite National Park (Miguel Meadow, 5,200' alt.), VII-10-38, on Pinus ponderosa (E. O. Essig, CAS). Shasta Co.: Lassen National Park, VI-19-35, on Pinus jeffreyi (W. H. Lange, CAS); 5 miles north Old Station, V-28-60, on Pinus ponderosa (H. H. Keifer, CDA).

Puto mimicus McKenzie, new species Mimic Mealybug (Figure 140) Type Locality and Host. Type and paratype adult females collected 5 miles north Truckhaven, Imperial County, California, on Encelia jrutescens (Compositae), June 10, 1963, by D. R. Miller. Additional paratype adult females collected on sandpaper plant, Petalonyx thurberi (Loasaceae), 6 miles south Valerie Jean, Riverside County, California, June 10, 1963, by D. R. Miller. Type Material. Holotype adult female (single specimen on slide) and female paratypes in museum, University of California, Davis. Paratypes in United States National Collection of Coccoidea, Washington, D.C., and in collection of California State Department of Agriculture, Sacramento. North American Distribution. Known only from California. Additional Hosts. None External Features and Habitat. This mealybug is

covered with a white, waxy secretion which is heavy and floculent. There are many small filaments along sides of body, some matted together, caudal filaments elongate, about three times as long as adjoining ones. The body color, which may be seen only in rubbed specimens, is dark green, and the legs are red. No ovisac was observed. The mealybug occurs on the foliage and stems of its host. Recognition Characters. Adult females quite small for members of this genus, mounted, 2.00 to 2.90 mm long; 1.20 to 2.00 mm wide; body form broadly oval. Dorsum basically with 18 pairs of cerarii, this arrangement, however, interrupted by division of individual cerarii or interpolation of others; each cerarius formed upon a sclerotized plate bearing from 2 or 3 enlarged, apically pointed and slightly lanceolate setae in smaller cerarii, to as many as 28 in anal lobe cerarii, or perhaps even more, and almost as many in other larger cerarii; without tubular ducts; trilocular pores borne on cerarian plates quite large, usually somewhat more numerous than enlarged setae. Trilocular pores quite large, of 2 different sizes, distributed rather evenly over entire dorsum. Minute circular pores scattered over body surface. Tubular ducts absent from dorsal body surface. Body setae for most part definitely smaller and more slender than those of cerarii, arranged in irregular rows across abdominal segments, scattered over thorax and head; few setae as large as or only slightly smaller than cerarian setae, situated in midregion and lateral area of most abdominal segments. Anal ring apical, with no unusual features; each of its 6 setae nearly twice as long as greatest diameter of ring. Venter with multilocular disk pores absent, replaced with what is here interpreted as modified quinquelocular pores, each with delicate outer ring, within which is a circle of small openings, and within this a small central ring divided into 5 cells, these pores absent on abdomen, sparingly distributed in sternal region, apparently absent on head. Trilocular pores sparingly distributed over venter, certain "clear" areas showing in sternal region where they apparently are absent. Very few oral-collar tubular ducts on abdomen and metathorax, absent anterior to mouthparts on head. Body setae few, for most part short and slender, those in midregion of abdomen noticeably longer. Circulus quite small, transversely oval. Legs unusually large and slender; hind tibia with translucent pores along entire segment; claw with prominent denticle or tooth on plantar surface. Antennae normally 9-segmented. Notes This species is a companion one with Puto nulliporus McKenzie, but differs in total absence of tubular ducts of any kind on dorsal surface, and more numerous, often as many as 28 or 30, enlarged conical setae in anal lobe cerarius. P. nulliporus, on the other hand, possesses oral-rim tubular type ducts, sparingly distributed but always present, on dorsum, and fewer, usually not more than 18, enlarged conical

TAXONOMY

OF

SPECIES

Fig. 140. Puto mimicus McKenzie, new species, collected on Encelia frutescens (Compositae), north of Truckhaven, Imperial County, California.

353

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setae in anal lobe cerarius. Their close similarity suggested the name from Latin mimicus (Greek mimikos), meaning imitative. California Records (Map 130) Imperial Co.: 5 miles north Truckhaven, VI-1063, on Encelia frutescens (D. R. Miller, UCD). Riverside Co.: 6 miles south Valerie Jean, VI10-63, on Petalonyx thurberi (D. R. Miller, UCD).

Puto nulliporus

McKenzie

Nulliporus Mealybug

(Figure 141) Puto nulliporus McKenzie, 1960. Hilgardia 29(15): 733-735, illus. Type Locality and Host. 9 miles south Palo Verde, Imperial County, California, on Bebbia juncea var. asper (Compositae). North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. No available information. Recognition Characters. Adult females, mounted, 2.50 to 3.50 mm long, 1.80 to 2.60 mm wide; body form broadly oval. Dorsum basically with 18 pairs of cerarii, this arrangement, however, interrupted by division of individual cerarii or interpolation of others; each cerarius formed upon a sclerotized plate bearing from 2 to 3 enlarged, apically pointed and slightly lanceolate setae in smaller cerarii, to as many as 18 in anal lobe cerarii, or perhaps even more, and almost as many

in other larger cerarii; without tubular ducts; trilocular pores borne on cerarian plates quite large, usually somewhat more numerous than enlarged setae. Trilocular pores quite large, of 2 different sizes, distributed over entire dorsum. Minute circular pores scattered over body surface. Tubular ducts of oral-rim type sparingly distributed over dorsum. Body setae for most part definitely smaller and more slender than those of cerarii, arranged in a broken single row on abdominal segments, scattered over thorax and head. Anal ring apical with no unusual features; each of its 6 elongate setae nearly twice as long as greatest diameter of ring. Venter with multilocular disk pores atypical for members of this genus, each with delicate outer ring, within which is a circle of small openings, and within this a small central ring defining 5 cells, these pores absent on last 5 abdominal segments (occasionally a single pore observed on fifth segment), sparsely distributed from fourth segment forward to sternal region of thorax. Trilocular pores sparingly distributed over venter, certain "clear" areas evident especially in sternal region where they are absent. Minute circular pores scattered on venter. One or two tubular ducts resembling oral-rim type, smaller than those on dorsum, may be found on abdomen. Very few oralcollar tubular ducts on thorax, particularly in sternal area, present or absent anterior to mouthparts on head. Body setae for most part long and slender, very sparsely distributed on ventral body surface. Circulus quite small, transversely oval. Legs unusually large and slender; hind tibia with translucent pores along entire segment; claw with prominent denticle or tooth on plantar surface. Antennae normally 9-segmented, slender. Notes. This species is very closely related to Puto mimicus McKenzie, but differs in possessing oral-rim tubular type ducts, sparingly distributed but always present, on dorsum, and fewer, usually not more than 18, enlarged conical setae in anal lobe cerarius. P. mimicus, on the other hand, lacks dorsal tubular ducts, and there are many more, often as many as 28 or 30, enlarged conical setae in anal lobe cerarius. California Records (Map 131) Imperial Co.: 9 miles south Palo Verde, IV-8-59, on Bebbia juncea var. asper (G. G. Beevor and W. P. Cronk, CDA). Puto pacificus McKenzie, new species

Pacific Mealybug (Figure 142) Type Locality and Host. Type and paratype adult females on sagebrush, Artemisia pycnocephala (Compositae), and plantain, Plantago juncoides (Plantaginaceae), at Goat Rock, Sonoma State Park, Sonoma County, California, collected December 14, 1963, by D. R. Miller. Type Material. Holotype adult female (single specimen on slide) and female paratypes in museum, University of California, Davis. Paratypes in United

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Fig. 141. Puto nulliporus McKenzie, collected on Bebbia juncea variety asper (Compositae), south of Palo Verde, Imperial County, California.

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TAXONOMY

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357

generally more oval in shape, scattered over dorsum. Tubular ducts, similar to those near cerarii, situated in median area of abdominal segments 8 - 5 , absent elsewhere on dorsum. Body setae for most part slightly smaller than those of cerarii. Anal ring apical, normal for genus; each of its 6 setae nearly twice as long as greatest diameter of ring itself . Venter with numerous multilocular disk pores, these all of approximately 1 size, distributed from posterior apex of abdomen forward to and including head region. Trilocular pores rather evenly distributed except for certain "clear" areas in sternal region where they seem to be entirely absent. Modified minute circular or discoidal-type pores similar to those on dorsum, scattered over venter. Oral-collar tubular ducts, smaller than those on dorsum, fairly numerous across most abdominal segments, scattered on thorax; a few more slender, elongate tubular ducts situated in cephalic region anterior to mouthparts. Body setae long and slender for most part exceeding in length those on dorsum. Circulus quite small, broadly oval, margins quite smooth. Legs comparatively short for members of genus; claw with prominent denticle on plantar surface. Antennae normally 9-segmented. States National Collection of Coccoidea (U. S. National M u s e u m ) , Washington, D. C., and in collection of California State Department of Agriculture, Sacramento. North American Distribution. California and Washington—apparently a coastal species. Additional Hosts. Gaultheria shallon (Ericaceae) and Vactinium sp. (Ericaceae). External Features and Habitat. This mealybug is heavily dusted with a white, fluffy, wax secretion. The body form is oval to nearly circular; body color pink, when crushed light brown. There are long broad waxy filaments around entire body, those at posterior extremity of body longest, shortest ones on head. No ovisac was observed. The male is wingless. T h e mealybug occurs on the crown and roots of its host and was tended by ants at the time of collection. Recognition Characters. Adult females not especially large for members of this genus, when mounted ranging from 2.60 to 3.50 mm long, 190 to 2.40 mm wide; body form broadly oval. Dorsum with basically 18 or 19 pairs of comparatively small sclerotized cerarian plates, this arrangement, however, interrupted by division of individual cerarii or interpolation of others; each cerarius formed upon a sclerotized plate bearing from 2 to 19 enlarged setae, or perhaps even more, these apically pointed, slightly lanceolate, trilocular pores present in numbers about equal to setae; 1 - 3 enlarged oral-collar type tubular ducts near most abdominal cerarii, these ducts with orifices slightly greater in diameter than diameter of socket at base of cerarian seta. Trilocular pores fairly evenly distributed on dorsum. Modified minute circular or discoidal-type pores,

Notes. This species shares with Puto cupressi ( C o l e m a n ) , the presence of dorsal enlarged oralcollar type tubular ducts along midregion of most abdominal segments, these in addition to those situated near most abdominal cerarii. It differs from P. cupressi in the absence of enlarged tubular ducts within the sclerotized margin of most cerarian plates; circulus smaller, short, broadly oval; only 6 enlarged anal ring setae; presence of a wingless male; and a much different host (Compositae and Plantaginaceae). On the other hand, P. cupressi possesses dorsal enlarged oral-rim type tubular ducts within margin of most cerarian plates; circulus transversely elongate oval, margins crenulated; anal ring frequently with 8 or more enlarged setae; a winged male; and normally occurs on Pinaceae. California Records (Map 132) Sonoma Co.: Goat Rock, Sonoma State Park, X I I - 1 4 - 6 3 , on Artemisia pycnocephala and Plantago juncoides (D. R. Miller, U C D ) .

Puto pricei McKenzie Price Mealybug (Figure 143) Puto pricei McKenzie, 1960. Hilgardia 2 9 ( 1 5 ) : 7 3 5 739, illus. Type Locality and Host. Yosemite National Park, Dana Fork of Tuolumne River (10,000 feet elevation), Tuolumne County, California, under rocks beneath lodgepole pine, Pinus contorta var. latifolia (Pinaceae). North American Distribution. Known only f r o m California. Additional Hosts. Pinus albicaulis (Pinaceae). External Features and Habitat. N o available infor-

358

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Fig. 143. Puto pricei McKenzie, collected under rocks beneath Lodgepole pine, Pinus contorta variety latifolia (Pinaceae), at Yosemite National Park, Dana Fork of Tuolumne River (10,000 feet elevation), Tuolumne County, California.

TAXONOMY

M a p 132.

Distribution

in California of Puto pacificus McKenzie.

mation on external appearance of this mealybug when alive. According to D. W. Price, who first collected the species, this mealybug was found at high altitudes of from 10,000 to 11,000 feet elevation, under and adhering tightly to rocks, and in needle debris around the base of lodgepole pine. He also found the mealybug under fallen logs beneath white bark pine. Partially grown nymphs and fully mature adult females were evident at the time these collections were made (September 6, 1959). No very definite ovisac is formed. The mealybugs apparently were not feeding on plant roots at the time collected. Recognition Characters. Adult females, mounted, 3.00 to 4.00 mm long, 1.80 to 2.80 mm wide; body form broadly oval. Dorsum basically with at least 18 pairs of cerarii, this arrangement, however, interrupted by division of individual cerarii or interpolation of others; each cerarius formed upon a sclerotized plate bearing from 3 or 4 to as many as 19 to 20 enlarged, apically pointed slightly lanceolate setae; without tubular ducts; trilocular pores borne on cerarian plates usually somewhat more numerous than the conical setae. Trilocular pores distributed over entire dorsum. Dorsal body setae similar in form to those of cerarii, numerous, arranged in transverse bands on abdominal segments, and covering thoracic segments and head. Anal ring apical, normal for genus; each of its 6 setae nearly twice as long as greatest diameter of ring. Venter with multilocular disk pores on last 2 abdominal segments, sparsely distributed in midregion from seventh segment forward to sternal region and head, somewhat clustered in coxal and spiracular areas, those anterior to last 2 abdominal segments be-

OF

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359

ing slightly smaller in diameter. Trilocular pores sparingly distributed over venter with exception of certain "clear" areas on mid-thoracic region where they appear to be entirely absent. Oral-collar tubular ducts lightly scattered over abdomen, 1 or 2 ducts situated anterior to mouthparts. Body setae long and slender, for most part exceeding in length those on dorsum. Circulus broad, transversely oval, margins appearing somewhat crenulated. Legs unusually large and slender; hind femur and tibia with translucent pores; claw with well-developed, prominent denticle or tooth on plantar surface, and a pair of rather unusual, sclerotized spurs at base. Antennae normally 9-segmented, slender. Notes. On the basis of high altitude habitat, this species would approach Puto bryanthi Ferris, and on the basis of an elongate, transversely oval circulus, it would come close to P. ambiguus (Fullaway). Among other things, it differs from P. bryanthi in the circulus character, and from P. ambiguus in possessing fewer ventral multilocular disk pores on abdominal segments 3 to 7 and in sternal and head areas. It may be distinguished from both P. bryanthi and P. ambiguus in having fewer, 20 or less, enlarged setae on anal lobe cerarius, as compared to 28 or more in the last 2 species mentioned. California Records (Map 133) Fresno Co.: 19 miles southwest Bishop (Upper Golden Trout Lake), IX-23-65, on Pinus albicaulis (bark crevices, foliage, roots) (G. M. Buxton, M. S. Wasbauer, and F. L. Blanc, C D A ) ; Mt. Goethe, V I I 9-52, on Pinus albicaulis (P. Raven, CAS). Tuolumne Co.: Yosemite National Park, Dana Fork Tuolumne River (10,000 feet elev.), IX-6-59, under rocks beneath Pinus contorta var. latifolia (D. W. Price, U C D ) .

M a p 133. Distribution in California of Puto pricei McKenzie.

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Puto profusus McKenzie Douglas Fir Mealybug (Figure 144) Puto profusus McKenzie, 1960. Hilgardia 29(15):738-741, illus. Type Locality and Host. Carrville, Trinity County, California, on Douglas fir, Pseudotsuga taxifolia (Pinaceae). North American Distribution. Known only from California. Additional Hosts. Abies sp. (Pinaceae). External Features and Habitat. No available information on appearance in life. It is suspected that this mealybug was collected from the foliage of its host. Recognition Characters. Adult females, mounted, 4.10 to 5.75 mm long, 2.75 to 3.80 mm wide; body form broadly oval. Dorsum basically with 18 pairs of cerarii, this arrangement, however, interrupted by division of individual cerarii or interpolation of others; each cerarius formed upon a sclerotized plate bearing from 2 or 3 to as many as 24, or perhaps even more, enlarged, apically pointed, slightly lanceolate setae; entirely without tubular ducts; trilocular pores in numbers equal to number of enlarged setae. Trilocular pores fairly evenly distributed over entire dorsum. Dorsal body setae as large as or only slightly smaller than cerarian setae, quite numerous, arranged in transverse bands on abdominal segments, more irregularly situated on thoracic segments and head. Anal ring apical, normal for genus; each of its 8 elongate setae nearly twice as long as greatest diameter of ring. Venter with comparatively few multilocular disk pores, all of same size, apparently restricted to abdominal segments 8 and 9, absent elsewhere on venter, except for 1 or 2 near thoracic spiracles. Trilocular pores distributed, mostly in bands, on abdomen and thorax, scattered on head. Very few oral-collar tubular ducts on abdominal segments; 1 or 2 more elongate ones situated in cephalic region anterior to mouthparts. Body setae long and slender, for most part exceeding in length those on dorsum. Circulus transversely elongate-oval, its margin appearing crenulated. Legs unusually large and slender; hind femur and tibia with translucent pores; claw with prominent denticle or tooth on plantar surface. Antennae normally 9-segmented, slender. Notes. One of the principal morphological differences of this mealybug, compared to other North American species of this group, is the absence of ventral multilocular disk pores in the thoracic and head regions, except for 1 or 2 near spiracles, the remainder confined to abdominal segments 8 and 9. It shares with P. cupressi (Coleman) a similar host preference and a transversely elongate-oval circulus with crenulated margin, but differs in the absence of tubular ducts associated with cerarian plates, these present in the species last mentioned.

California Records (Map 134) Plumas Co.: 1 mile south Prattville, VII-9-64, on Abies sp. (under bark of trunk, less than 4 feet high) (D. R. Miller, UCD). Trinity Co.: Carrville, VI-1933, on Pseudotsuga taxifolia (E. C. Van Dyke, CDA).

Puto simmondsiae McKenzie Simmondsia Mealybug (Figure 145) simmondsiae McKenzie, 1961. Hilgardia 31(2) :39—41, illus. Type Locality and Host. Indio, Riverside County, California, on goat-nut or Jajoba, Simmondsia chinensis (Buxaceae). North American Distribution. California and Mexico. Additional Hosts. None. External Features and Habitat. No available information, except that the mealybug occurs above ground on the leaves and stems of its host. Recognition Characters. Adult females, mounted, 3.20 to 3.80 mm long, 2.00 to 2.20 mm wide; body form broadly oval. Dorsum basically with 18 pairs of cerarii, this arrangement, however, interrupted by division of individual cerarii or by interpolation of others; each cerarius formed upon a sclerotized plate with 2 to 5 tubular ducts of oral-rim type, these possibly somewhat variable in size; 11 to 28, or perhaps more, enlarged, conical, apically pointed, slightly lanceolate setae; trilocular pores usually somewhat more numerous than conical setae.

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Fig. 144. Puto profusus McKenzie, collected on Douglas fir, Pseudotsuga (Pinaceae), at Carrville, Trinity County, California.

taxifolia

361

362

TAXONOMY

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Fig. 145. Puto simmondsiae McKenzie, collected on Jajoba or goat-nut, chinensis (Buxaceae), at Indio, Riverside County, California.

Simmondsia

TAXONOMY Trilocular pores evenly beset over entire dorsum. Minute circular pores, somewhat sclerotized, scattered over dorsal surface. Tubular ducts, of type represented within cerarian plates, sparingly distributed on thorax. Body setae, similar to those of cerarii, numerous and arranged in transverse bands on abdominal segments and covering thoracic segments and head. Anal ring apical, normal for genus; each of its 6 setae nearly twice as long as greatest diameter of ring. Venter with multilocular disk pores situated mainly in midregion of last 3 abdominal segments, sparsely distributed near spiracles, front coxae, mouthparts, absent on head. Trilocular pores sparingly distributed over venter, except for certain "clear" areas in sternal region where they appear entirely absent. Minute circular pores, somewhat sclerotized, scattered over ventral surface. Tubular ducts of oral-collar type situated, in limited numbers, on venter of abdominal segments; few more slender ducts noticeable in cephalic region anterior to mouthparts. Body setae long and slender, for most part exceeding in length those on dorsum. Circulus broad, transversely oval, margins appearing somewhat crenulated. Legs unusually large; claw with prominent denticle or tooth on plantar surface. Antennae normally 9-segmented, slender. Notes. This species is similar to Puto yuccae (Coquillett), particularly in restricted number of tubular ducts on cerarian plates. It differs from P. yuccae, however, in lacking ventral multilocular disk pores on abdomen anterior to sixth segment, and far fewer of these structures on thorax, absent on head. P. yuccae, on the other hand, possesses numerous ventral multiloculars on all abdominal segments, thorax, and head. California Records (Map 135) Riverside Co.: Indio, IX-9-35, on Simmondsia chinensis (H. L. McKenzie, UCD). Silkiifoi, Tfinitu / SmistaLassen Jifama Pluma»

Map 135.

Distribution

in California of Puto . simmondsiae McKenzie.

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Puto yuccae (Coquillett) Large Yucca Mealybug (Figure 146; Color Plate XX) Dactylopius crawii Coquillett, 1889. West Amer. Sci. 6:123 (as to the male). Dactylopius yuccae Coquillett, 1890. Ibid. 7:44. Phenacoccus yuccae (Coquillett), Cockerell, 1896. 111. State Lab. Nat. Hist. 4:318-339. Phenacoccus bahiae Ehrhorn, 1900. Canad. Ent. 32:315. Ceroputo yuccae (Coquillett), Cockerell, 1901. Proc. Biol. Soc. Wash. 14:166. Ceroputo bahiae (Ehrhorn), Cockerell, 1901. Canad. Ent. 33:166. Phenacoccus simplex King, 1902. Ent. News 13:42-43. Ceroputo yuccae var. ceanothi Cockerell, 1903. Ann. Mag. Nat. Hist. (ser. 7) 11:163. Phenacoccus ramonae Essig, 1909. Pomona Col. Jour. Ent. 1(2):44, illus. Ceroputo yuccae (Coquillett), Essig, 1909. Ibid. 1(4):94-95, illus. Ceroputo yuccae (Coquillett), Essig, 1914. Calif. State Comn. Hort. Monthly Bull. 3 ( 3 ) : 1 2 3 124, illus. Puto yuccae (Coquillett), Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., pp. 64-65. Puto yuccae (Coquillett), Ferris, 1950. Atlas of scale insects of North America, Ser. V:213-215, illus. Puto yuccae (Coquillett), Ferris, 1953. Ibid., Vol. VI: 423. Puto yuccae (Coquillett), McKenzie, 1961. Hilgardia 31(2):32-33 (key). .Type Locality and Host. Los Angeles, Los Angeles County, California, on Our Lord's Candle, Yucca whipplei (Agavaceae). North American Distribution. California, Mexico, and Texas. Additional Hosts. It is quite evident that this species has an extremely wide range of host plants. Some are as follows: Arctostaphylos spp. (Ericaceae), Astragalus miguelensis (Leguminosae), Atriplex sp. (Chenopodiaceae), Baccharis viminea (Compositae), Batis ? sp. (Batidaceae), Ceanothus oliganthus (Rhamnaceae), Citrus limon (Rutaceae), Eriophyllum confertiflorum (Compositae), Garrya elliptica, G. veatchii (Garryaceae), Lepidospartum squamatum (Compositae), Mesembryanthemum sp. (Aiozoaceae), Oenothera sp. (Onagraceae), Salvia (= Ramona) polystachya (Labiatae), Sebastiania bilocularis (Euphorbiaceae), and Stachys bullata (Labiatae). External Features and Habitat. The adult female of this species is thickly covered with white, waxy secretion, equipped with short, stout lateral filaments and somewhat longer caudal ones. An ovisac is not formed (see col. pi. XX). The adult male is winged, with a wing expanse of 7.00 to 8.00 mm, and a pair of long caudal wax filaments measuring as much as 6.00 mm. The mealybug occurs mainly on the stems and foli-

364

TAXONOMY

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Fig. 146. Puto yuccae (Coquillett), collected on Our Lord's Candle, Yucca (Agavaceae), at Los Angeles, Los Angeles County, California.

whipplei

COLOR PLATE XX. Large yucca mealybug, Puto yuccae (Coquillett), on leaves of veatch silk tassel, Garrya veatchii (Garryaceae), near Cajon, San Bernardino County, California. Note winged male on stem (right), and male pupal case on uppermost leaf.

age of its host, although at times is found on the roots. Recognition Characters. Adult female, mounted, 3.00 to 5.00 mm long, 2.00 to 3.25 mm wide; body form broadly oval. Dorsum basically with 18 pairs of cerarii, this arrangement, however, interrupted by division of individual cerarii or interpolation of others; each cerarius formed upon moderately large, circular or oval, sclerotized plate bearing from usually 6 to 8 to as many as 30, or perhaps even more, enlarged setae, these apically pointed, slightly lanceolate, also bearing 2-4 tubular ducts—number quite constant, most frequently 2 and rather rarely as many as 4—these quite variable, but typically slightly larger in diameter than diameter of basal socket of cerarian setae, and trilocular pores usually almost as numerous as enlarged setae. Trilocular pores fairly evenly distributed on dorsum. Discoidal type pores lightly scattered on dorsal surface. Dorsal body setae for most part slightly smaller than cerarian setae, each with very slight basal tubercle—some lightly sclerotized—more or less scattered, and showing little or no tendency to be aggregated into groups. Anal ring apical, normal for genus; each of its 6 setae nearly twice as long as greatest diameter of ring. Venter with multilocular disk pores of 2 slightly different sizes, the smaller size situated on anterior portion of abdomen, forward to thoracic and cephalic regions. Trilocular pores sparingly distributed over venter, certain "clear" areas evident in sternal region. Discoidal type pores lightly scattered on ventral surface. Oral-collar tubular type ducts situated along posterior margin of most abdominal segments; several more slender and elongated ones noticeable in head area anterior to mouthparts. Body setae slender, for most part exceeding in length those on dorsum. Circulus transversely elongate oval, with crenulate margin. Legs comparatively large; claw with prominent denticle or tooth on plantar surface. Antennae normally 9-segmented. Notes. This species is similar to Puto simmondsiae McKenzie, particularly in restricted number of tubular ducts on cerarian plates. It differs from P. simmondsiae in possessing numerous ventral multiloculardisk pores on all abdominal segments, thorax, and head, whereas in the species last mentioned, these structures are absent on abdomen anterior to sixth segment, far fewer of them on thorax and absent on head. California Record (Map 136) Glenn Co.: Alder Springs, VII-7-33, on Arctostaphylos sp. (H. H. Keifer, CD A). Inyo Co.: Between Bishop and Mammoth, VII1918, on Oenothera sp. (G. F. Ferris, UCD). Lake Co.: Lower Lake, V-25-59, on Eriodictyon californicum (T. R. Haig, CDA). Los Angeles Co.: Azusa, VII-6-39, on Lepidospartum squamatum (L. E. Myers, LEM); 2 miles north Azusa, San Gabriel Canyon, VII-29-62, on Batis ? sp. (W. D. Dyer, CDA); El Monte, X-25-

Map 136. Distribution in California of Puto yuccae (Coquillett).

Smümcif

38, on ? (K. E. Frick, UCD); 2 miles south Kentucky Springs, VI-11-63, on Yucca sp. (L. E. Myers, CDA); 15 miles west Littlerock (Little Rock Creek), VII-25-63, on Yucca whipplei (leaves) (D. R. Miller and D. R. Estes, UCD) ; Long Beach, VII-20-23, on Baccharis viminea (E. Bethel, CDA) ; San Dimas, V-5-16, on Citrus sp. (orange) ( W. E. Landon, CAS) (immature) ; Saugus, XII-20-16, on ? (R. W. Doane, CDA); Saugus (Solidad Canyon), 1939, on Eriogonum jasciculatum (L. E. Myers, LEM). Napa Co.: 5 miles northeast of Napa, VIII-3-63, on Diplacus aurantiacus (roots) (D. R. Miller, UCD). Orange Co.: Anaheim, 1910, on "sage" (R. W. Doane, CDA); 2 miles north Modjeska Canyon, VI-7-63, on Diplacus sp. (roots, crowns, and foliage) and Eriophyllum confertiflorum (leaves) (D. R. Miller, UCD). Riverside Co.: Idyllwild, VIII-5-33, on Arctostaphylos sp. (A. J. Bassinger, UCD); San Jacinto Mts., VIII-1938, on ? (Beamer, UCD). San Bernardino Co. : 2 miles south Cajon, VI-2464, on Garrya veatchii (roots and foliage) (D. R. Miller, UCD). San Diego Co.: Palomar Mts., VII-11-51, on Quercus dumosa (F. L. Blanc, CDA); Tecata (Peak), VI-15-63, on Chamaebatia australis (B. C. Reed, CDA); Tecate, VI-2-59, on Chamaebatia foliolosa (S. Klopfer, CDA). San Francisco Co.: San Francisco (sand hills), III-1913, on Mesembryanthemum sp. (T. C. Bridwell, CDA). San Luis Obispo Co.: San Miguel, VII-1937, on Astragalus miguelensis (T. D. A. Cockerell, CDA). Santa Barbara Co.: Lompoc (Vandenberg Village), VI-5-63, on Eriastrum densifolium, Horkelia

366

TAXONOMY

OF

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sp., and Salvia columbariae (roots) (D. R. Miller, UCD). Santa Clara Co.: Stanford University, V-3-17, on Stachys bullata (G. F. Ferris, U C D ) ; Stevens Creek, X-14-16, on Eriophyllum confertiflorum (G. F. Ferris, U C D ) .

PYGMAEOCOCCUS Pygmaeococeus McKenzie, 1960. Hilgardia, 2 9 ( 1 5 ) : 741. Genotype. Pygmaeococcus morrisoni McKenzie. Recognition Characters. Extremely small Pseudococcidae, adult female, at maturity, measuring not more than 0.75 mm in length, 0.20 mm in width. Dorsal ostioles well developed although not sclerotized. Dorsum with cerarii entirely absent; anal lobes commonly with several long setae. Trilocular pores present on both surfaces, these comparatively large. Bi- or tritubular pores and multilocular disk pores absent. Dorsum and venter with unusual type invaginated tubular ducts, with what appears to be a septum across inner end. Small, short, slender setae sparsely distributed on both body surfaces. Anal ring apical, relatively small, but well developed and complete, its pores large, oval or irregularly shaped; each of its 6 setae nearly twice as long as greatest diameter of ring. Circulus small, almost circular, situated on fourth abdominal segment. Legs stout, usually rather conspicuously spinose because of development of enlarged setae on inner margin of tibia and tarsus; claw without denticle on plantar surface; digitules equaling or slightly exceeding claw in length, apically swollen. Eyes present, well developed. Antennae 5segmented, set close together at apex of head, their bases ordinarily separated by less than width of basal segment, sensory setae of terminal segment strongly developed, unusually long and quite large. Notes. This is a most unusual mealybug, distinguishable from other known pseudococcids by its extremely small size at maturity, averaging 0.50 mm in length, 0.20 mm in width, and by the possession of rather remarkable invaginated tubular ducts present on both surfaces of body. It is perhaps most closely related to Rhizoccus Kunckel d'Herculais, differing principally in the characters just mentioned.

Pygmaeococcus morrisoni McKenzie Morrison Pygmy Mealybug (Figure 147) Pygmaeococcus morrisoni McKenzie, 1960. Hilgardia 2 9 ( 1 5 ) : 7 4 0 - 7 4 2 , illus. Type Locality and Host. Coalinga, 5 miles west in Warthon Canyon, Fresno County, California, collected in leaf mold beneath Juniper, Jurtiperus sp. (Cupressaceae).

Ventura Co.: Fillmore, IV-25-34, on Citrus limon (roots) (E. L. Smith, CD A ) ; Santa Paula, date ?, on Ramona stachyoides (roots) (E. O. Essig, CAS); 2 miles west Ventura, VIII-24-47, on Mesembryanthemum sp. (J. B. Steinweden, C D A ) .

MCKENZIE North American Distribution. Known only from California. Additional Hosts. "Bunch grass" (Gramineae). External Features and Habitat. No available information on external appearance of this mealybug in life. The mealybug has been found in leaf mold, and in soil on roots of "bunch grass." The specimens in leaf mold were recovered using a Berlese funnel. Recognition Characters. Adult females, mounted, average 0.50 mm long, 0.20 mm wide; body form elongate, sides almost parallel. Dorsum with cerarii entirely absent; anal lobes commonly with several long setae. Trilocular pores comparatively large, internal area of each quite irregular, pores sparsely scattered over dorsal surface. With unusual type invaginated tubular ducts, with what appears to be a septum across inner end, sparsely scattered over dorsal surface. Body setae small, slender, lightly distributed over body. Anal ring apical, comparatively small, but well developed and complete, its pores large, oval or irregularly shaped; each of its 6 setae nearly twice as long as greatest diameter of ring. Venter without multilocular disk pores. Trilocular pores similar to those on dorsum, sparsely distributed over venter. Invaginated tubular ducts, similar in size and shape to those on dorsum, sparsely distributed. Body setae short and slender. Circulus small, almost circular, situated on fourth abdominal segment (1 specimen shows 2 circuli). Legs stout, usually rather conspicuously spinose because of development of enlarged setae on inner margin of tibia and tarsus; claw without denticle on plantar surface; digitules equaling or slightly exceeding claw in length, apically swollen. Antennae 5-segmented, set close together at apex of head, their bases ordinarily separated by less than width of basal segment, sensory setae of terminal segment strongly developed, unusually long and quite large. Notes. This is a most unusual mealybug, distinguishable from other known pseudococcids by its extremely small size, averaging 0.50 mm in length, 0.20 mm in width, at maturity, and in the possession of rather remarkable invaginated tubular ducts situated on both surfaces of the body. It is perhaps most closely related to certain Rhizoecus species, differing principally in the characters just mentioned. A single specimen collected on roots of a bunch grass shows 2 circuli instead of only 1 although all

TAXONOMY

OF

SPECIES

Fig. 147. Pygmaeococcus morrisoni McKenzie, collected on "bunch grass" (root) (Gramineae), north of Hat Creek, Shasta County, California. (Illustration prepared from type material.)

367

368

TAXONOMY

OF

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other features agree in detail with the type lot, and it is suspected this is a normal variation. California Records (Map 137) Fresno Co.: Coalinga, 5 miles west in Warthon Canyon, VI-25-58, in "juniper leaf mold" (H. L. Wilson, C D A ) . Shasta Co.: 10.7 miles north Hat Creek, V - 2 3 - 6 3 , "bunch grass" (root) (T. R. Haig, C D A ) .

RADICOCCUS Radicoccus Hambleton, 1946. Rev. de Ent. 1 7 ( 1 2):47. Radicoccus Hambleton, Ferris, 1953. Atlas of the scale insects of North America, Vol. VI: 424. Genotype. Rhizoecus globosus James. Recognition Characters. As based upon the single North American species referred to this genus, Radicoccus may be described as follows: Pseudococcidae with dorsal ostioles absent or at least extremely obscure. Cerarii absent. Pores and ducts very few, these including merely a few small trilocular pores and a few very short, tubular ducts. Body setae almost completely lacking, there being apparently only 2 pairs marking position of anal lobes, absent elsewhere on body. Anal ring borne on dorsal surface, well removed from posterior apex of abdomen, ring complete, very few relatively large pores; each of its 6 setae short, not equaling in length diameter of ring itself. Two small, circular circuli situated medially on fourth and fifth abdominal segments. Legs very small, but completely developed; claw without denticle on plantar surface; digitules knobbed. Antennae 5-segmented, short, stout, set quite close together at their bases. Spiracles extremely small. Eyes absent. Notes. The total absence of setae anywhere on the body, except for 2 pairs which mark position of the anal lobes; and the comparatively very small legs and antennae, distinguish this group from other California genera. De Lotto (1964) proposed that Radicoccus Hambeton be placed as a synonym of Rhizoecus Kiinckel d'Herculais. I am inclined to retain Radiococcus as a valid genus mainly on the basis of the very peculiar

HAMBLETON type of tubular ducts with elliptical orifices, the almost total absence of body setae, the extremely small spiracles, and the very much reduced legs and antennae.

Radicoccus kelloggi (Ehrhorn and Cockerell) Kellogg Mealybug (Figure 148) Ripersiella kelloggi Ehrhorn and Cockerell, Cockerell, 1901. Proc. Biol. Soc. Wash. 14:165-166. Ripersiella kelloggi Ehrhorn and Cockerell, Ehrhorn, 1906. Canad. Ent. 3 8 ( 1 0 ) : 3 3 4 . Ripersiella kelloggi Ehrhorn and Cockerell, Fernald, 1903. Mass. Agr. Expt. Sta. Spec. Bull. 88, p. 115. Ripersiella kelloggi Ehrhorn and Cockerell, Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., p. 73, illus. Ripersiella kelloggi Ehrhorn and Cockerell, Cockerell and Bueker, 1930. Am. Mus. Nat. Hist.,'Amer. Mus. Novitates 441:1. Radicoccus kelloggi (Ehrhorn and Cockerell), Hambleton, 1946. Rev. de Ent. 1 7 ( 1 - 2 ) : 5 0 , illus. Radicoccus kelloggi (Ehrhorn and Cockerell), Ferris, 1953. Atlas of scale insects of North America, Vol. VI: 424-425, illus. Type Locality and Host. Stevens Creek, near Mountain View, Santa Clara County, California, on "bunch grass" (Gramineae). North American Distribution. California, Connecticut, and New Jersey. Additional Hosts. Ammophila breviligulata (Gra-

368

TAXONOMY

OF

SPECIES

other features agree in detail with the type lot, and it is suspected this is a normal variation. California Records (Map 137) Fresno Co.: Coalinga, 5 miles west in Warthon Canyon, VI-25-58, in "juniper leaf mold" (H. L. Wilson, C D A ) . Shasta Co.: 10.7 miles north Hat Creek, V - 2 3 - 6 3 , "bunch grass" (root) (T. R. Haig, C D A ) .

RADICOCCUS Radicoccus Hambleton, 1946. Rev. de Ent. 1 7 ( 1 2):47. Radicoccus Hambleton, Ferris, 1953. Atlas of the scale insects of North America, Vol. VI: 424. Genotype. Rhizoecus globosus James. Recognition Characters. As based upon the single North American species referred to this genus, Radicoccus may be described as follows: Pseudococcidae with dorsal ostioles absent or at least extremely obscure. Cerarii absent. Pores and ducts very few, these including merely a few small trilocular pores and a few very short, tubular ducts. Body setae almost completely lacking, there being apparently only 2 pairs marking position of anal lobes, absent elsewhere on body. Anal ring borne on dorsal surface, well removed from posterior apex of abdomen, ring complete, very few relatively large pores; each of its 6 setae short, not equaling in length diameter of ring itself. Two small, circular circuli situated medially on fourth and fifth abdominal segments. Legs very small, but completely developed; claw without denticle on plantar surface; digitules knobbed. Antennae 5-segmented, short, stout, set quite close together at their bases. Spiracles extremely small. Eyes absent. Notes. The total absence of setae anywhere on the body, except for 2 pairs which mark position of the anal lobes; and the comparatively very small legs and antennae, distinguish this group from other California genera. De Lotto (1964) proposed that Radicoccus Hambeton be placed as a synonym of Rhizoecus Kiinckel d'Herculais. I am inclined to retain Radiococcus as a valid genus mainly on the basis of the very peculiar

HAMBLETON type of tubular ducts with elliptical orifices, the almost total absence of body setae, the extremely small spiracles, and the very much reduced legs and antennae.

Radicoccus kelloggi (Ehrhorn and Cockerell) Kellogg Mealybug (Figure 148) Ripersiella kelloggi Ehrhorn and Cockerell, Cockerell, 1901. Proc. Biol. Soc. Wash. 14:165-166. Ripersiella kelloggi Ehrhorn and Cockerell, Ehrhorn, 1906. Canad. Ent. 3 8 ( 1 0 ) : 3 3 4 . Ripersiella kelloggi Ehrhorn and Cockerell, Fernald, 1903. Mass. Agr. Expt. Sta. Spec. Bull. 88, p. 115. Ripersiella kelloggi Ehrhorn and Cockerell, Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., p. 73, illus. Ripersiella kelloggi Ehrhorn and Cockerell, Cockerell and Bueker, 1930. Am. Mus. Nat. Hist.,'Amer. Mus. Novitates 441:1. Radicoccus kelloggi (Ehrhorn and Cockerell), Hambleton, 1946. Rev. de Ent. 1 7 ( 1 - 2 ) : 5 0 , illus. Radicoccus kelloggi (Ehrhorn and Cockerell), Ferris, 1953. Atlas of scale insects of North America, Vol. VI: 424-425, illus. Type Locality and Host. Stevens Creek, near Mountain View, Santa Clara County, California, on "bunch grass" (Gramineae). North American Distribution. California, Connecticut, and New Jersey. Additional Hosts. Ammophila breviligulata (Gra-

TAXONOMY

(Gramineae), at Pacheco Pass, Santa Clara County, California.

OF

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369

370

TAXONOMY

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mineae), Festuca ovina (Gramineae), and Stipa sp. (Gramineae). External Features and Habitat. According to Ehrhorn ( 1 9 0 6 ) , the adult female of this species produces a small quantity of white, cottony secretion which generally encases the body. Its body is creamywhite, and its shape is broadly oval, sometimes pyriform. The mealybug was described as from the roots of "bunch grass." Ferris (1953) states that specimens he collected were attached to very small rootlets, at some distance from the base of the plant itself. Recognition Characters. Adult females, mounted, 1.10 to 2.00 mm long, 0.90 to 1.50 mm wide; body form almost circular or pyriform. Dorsum with cerarii completely absent. Trilocular pores absent. Few dorsal, small tubular ducts which open through an elliptical pore, the rim of which is strongly sclerotized, these ducts scarcely longer than their own diameter, difficult to recognize except in profile. There is faint suggestion that ducts are divided by median partition. Body setae sparse, there being apparently 2 pairs marking position of anal lobes, otherwise lightly scattered on dorsum. Anal ring borne on dorsal surface, well removed from posterior apex of abdomen, ring complete, very few relatively large pores; each of its 6 setae short, scarcely exceeding in length half the diameter of ring itself. Trilocular pores very sparsely distributed on abdomen and submarginally on thorax and head. Tubular ducts, similar in size and shape to those on dorsum, sparsely distributed on ventral surface. Two small, circular circuli situated medially on fourth and fifth abdominal segments. Legs very small though completely developed; claw without denticle on plantar surface; digitules knobbed. Antennae short, stout, 5-segmented, set quite close together at their bases. Spiracles extremely small. Eyes absent. Notes. The absence of body setae, except for 2 pairs marking the position of the anal lobes; and the very small legs and antennae separate this species quite readily from all other California pseudococcids.

RHIZOECUS

This is the only species of Radicoccus known to occur in North America. California Records (Map 138) Del Norte Co.: 5 miles west of Smith River, V I I I 24-63, "under rock" (T. R. Haig, C D A ) . Nevada Co.: Truckee, V I I I - 2 6 - 6 5 , on Gramineae (T. R. Haig, C D A ) . Santa Clara Co.: Pacheco Pass, 111-22-35, on Stipa sp. (G. F. Ferris, U C D ) ; Steven Creek, near Mountain View, X I I - 1 8 9 8 , on "bunch grass" (E. M. Ehrhorn, U C D ) . Siskiyou Co.: 2 miles north Mt. Shasta, X I - 9 - 6 5 , on Festuca ovina (T. R. Haig, C D A ) . Trinity Co.: 6 miles northeast Weaverville, V - 2 2 - 6 3 , on "unknown shrub" (R. F. Wilkey, CDA).

Map 138. Distribution in California of

Radicoccus kelloggi (Ehrhorn and Cockerell).

San

frartciteoSan Motto

KÜNCKEL D'HERCULAIS

Rhizoecus Künckel d'Herculais, 1878. Ann. Ent. Soc. de France (Ser. 5) 8:163. Morrisonella Hambleton, 1946. Rev. de Ent. 1 7 ( 1 2):16. Rhizoecus Künckel d'Herculais, Hambleton, 1946. Ibid. 1 7 ( 1 - 2 ) : 5 0 - 5 1 . Coccidella Hambleton, 1946. Proc. Biol. Soc. Wash. 59:177. Rhizoecus Künckel d'Herculais, Ferris, 1953. Atlas of the scale insects of North America, Vol. VI: 426-427. Rhizoecus Künckel d'Herculais, Williams, 1962. Bull. Brit. Mus. (Nat. Hist.) Ent. 1 2 ( 1 ) :41.

Rhizoecus Kunckel d'Herculais, De Lotto, 1964. Ibid. 14(8):381—382. Genotype. Rhizoecus falcifer Kunckel d'Herculais. Recognition Characters. Pseudococcidae with dorsal ostioles usually present, very commonly with slight sclerotization of their rims, at times very obscure, possibly even absent. Dorsal cerarii entirely absent, anal lobes commonly with several slender setae. Multilocular disk pores present or absent. In addition to the usual trilocular pores, there are ones which seem to be derived from trilocular type, but are more or less enlarged, strongly sclerotic, with

TAXONOMY walls of the loculi prolonged to form tubes which project from body wall, and at times somewhat similar pores having only 2 such tubes—these called "tritubular" and "bitubular" pores, respectively. Occasionally tubular ducts of very small size and oralcollar type present. Anal ring relatively small, but well developed and complete, its pores large, oval or irregularly polygonal. Circuli varying from 0 to 2, these normally very small and circular. Legs usually spinose because of strongly developed setae on inner margin of tibia and tarsus; claw without denticle on plantar surface; digitules at times very short, at times equaling claw in length, never apically swollen. Antennae 5- to 6-segmented, ordinarily strongly geniculate, set close together at apex of head, their bases ordinarily not separated by more than twice width of basal segment; sensory setae of terminal segment strongly developed and unusually large. Head region anterior to clypeus usually with small sclerotized plate on ventral side. Eyes present or absent. Notes. Hambleton (1946) studied in great detail all the hypogeic mealybugs belonging in Rhizoecus and related genera. Ferris (1953) presented what is believed to be a somewhat sounder generic appraisal of Rhizoecus, and his opinions are here accepted. Apparently, the absence of eyes, number of antennal segments, presence or absence of circuli, presence or absence of bi- or tritubular pores, and length of claw digitules do not in themselves have much significance as generic characters. It is suspected, however, that with the discovery and description of more of these obscure, soil-inhabiting species—and when they are better known—more generic segregates will have to be considered. The bi- or tritubular pores seem quite distinctive for Rhizoecus and this character alone seems to differentiate it from other pseudococcid genera. Two new California species of Rhizoecus are described in this book. To the genus there are now assigned 29 species for North America, 17 of which occur in California.

6(4).

7(6). 8(7).

9(2). 10(9).

11(10). 12(11).

13(12).

14(13).

15(13).

16(15).

KEY Species of Rhizoecus in North America: Adult Females

1. 2(1).

3(2). 4(3).

5(4).

Without a circulus 17 With at least 1 circulus 2 Tritubular pores definitely present, although at times quite small and inconspicuous 3 Tritubular pores definitely absent . . . . 9 Antennae 5-segmented . californicus Ferris Antennae 6-segmented 4 Tubular ducts of oral-collar type absent 5 With at least a few tubular ducts of oral-collar type present, usually very small 6 Multilocular disk pores absent ventrally on abdomen . . leucosomus (Cockerell)

17(1).

18(17).

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371

Multilocular disk pores present ventrally on abdomen . sonomae McKenzie Multilocular disk pores absent ventrally on abdomen 7 Multilocular disk pores present ventrally on abdomen . . brownï McKenzie With 2 circuli bicirculus McKenzie With but 1 circulus 8 Digitules long and slender, weakly knobbed, extending to tip of claw . . cacticans (Hambleton) Digitules short, setose, not extending beyond middle of claw maritimus (Cockerell) With 3 to 5 circuli . campèstris Hambleton With not more than 2 circuli 10 Multilocular disk pores present ventrally on thorax bituberculatus McKenzie Multilocular disk pores absent on thorax venter 11 With 2 circuli kondonis Kuwana With but 1 circulus 12 Entire eighth abdominal segment with 35 or more ventral multilocular disk pores spinosus McKenzie Entire eighth abdominal segment with 25 or fewer ventral multilocular disk pores 13 Tarsal claws with digitules as long as or longer than claw itself, digitules apically knobbed or swollen 14 Tarsal claws with digitules shorter than claw itself, apically acute 15 Bitubular pores present gracilis McKenzie Bitubular pores absent, unitubular-type pore present . . . . disjunctus McKenzie Multilocular disk pores present on dorsum of eighth abdominal segment . . 16 Multilocular disk pores absent on dorsum of eighth abdominal segment . . totonicopanus Hambleton With paired tubular pores, the openings of which are set close together, the ducts parallel or somewhat divergent solani Hambleton Bitubular pores seemingly formed of a sclerotized cone, from the apex of which arise 2 curving ducts or tubes menkei McKenzie Dorsum of most abdominal segments with a transverse row of tritubular pores, these rows having 10 to 16 pores; antennae 5-segmented falcifer Kunckel d'Herculais Dorsum of abdominal segments with tritubular pores, if tritubulars are present, not thus arranged or if rows can be detected, they include not more than 6 to 7 pores 18 Anal lobes terminate in a small but dis-

372

19(18). 20(19).

21(18).

22(21).

23(22). 24(23).

25(24).

26(21).

27(26).

28(27).

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SPECIES

tinct area of sclerotization 19 Anal lobes without such a sclerotization 21 Bitubular pores present somewhere on body boharti McKenzie Bitubular pores absent on body 20 Inner rows of pores of the anal ring clouded associatus (Hambleton) Inner row of pores of the anal ring not clouded distinctus (Hambleton) Multilocular disk pores present only on venter of abdomen 22 Multilocular disk pores—even if very few—present dorsally and ventrally 26 Abdominal segments 6 and 7 ventrally each with a transverse row of small tritubular pores extending across median region . . mayanus (Hambleton) No abdominal segment with transverse row of small tritubular pores in median region 23 Eyes absent eluminatus McKenzie Eyes present 24 A very few small ventral tubular ducts present posterior to vulva 25 Small ventral tubular ducts absent anywhere on body . . pritchardi McKenzie Each anal lobe area with 2 elongate setae and 2 or 3 smaller auxiliary setae; 1 to 4 body setae associated with each tritubular pore cyperalis (Hambleton) Each anal lobe area with 3 elongate setae and 5 to 8 smaller auxiliary setae; body setae not associated with tritubular pores nemoralis (Hambleton) Dorsal multilocular disk pores form distinct rows on all segments from mesothorax to abdominal segment eight graminis (Hambleton) Dorsal multilocular disk pores very few and scattered, not in rows 27 Entire venter of eighth abdominal segment with 18 or fewer, usually fewer, multilocular disk pores neomexicanus McKenzie Entire venter of eighth abdominal segment with 28 or more multilocular disk pores 28 Entire venter of seventh and eighth abdominal segments with 20 or more tritubular pores stangei McKenzie Entire venter of seventh and eighth abdominal segments with 10 or fewer tritubular pores americanus (Hambleton)

Rhizoecus associatus (Hambleton) Associated Mealybug (Figure 149) Morrisonella associata Hambleton, 1946. Rev. de Ent. 17(1-2) :20-21, illus. Rhizoecus associatus (Hambleton), Ferris, 1953. Atlas of scale insects of North America, Vol. VI: 430-431, illus. Type Locality and Host. On road between Quetzaltenango and Totonicapan, Guatemala (10,000 feet elevation) on roots of Alchemilla orbiculata (Rosaceae). North American Distribution. California and Guatemala. Additional Hosts Unidentified Gramineae. External Features and Habitat. This mealybug is covered with a white, waxy secretion which is very diverse and difficult to see. The body form is elongate; body color white; legs white; and there are no filaments. A filamentous ovisac is formed which is approximately one and one-half times the length of mature adult female. The mealybug was found feeding on grass roots and also distributed in the soil with apparently no root-host association. Recognition Characters. Adult females, mounted, 1.30 to 2.00 mm long, 0.60 to 1.30 mm wide; body form elliptical, narrower at each end; antennae set close together at apex of head. Dorsal ostioles weakly developed. Anal lobe area definitely sclerotized, bearing 4-7 elongate setae. Dorsum rather sparsely beset with trilocular pores. Multilocular disk pores scattered over almost entire dorsum. Tritubular pores rather sparse, distributed over dorsal surface. Tubular ducts absent. Body setae of varying length beset on dorsal surface. Anal ring relatively large, with a triple row of large, irregular pores, the inner row lightly sclerotized; its 6 setae slightly longer than diameter of ring itself. Ventral multilocular disk pores distributed similar to those on dorsum with perhaps more concentration about vulva. Trilocular pores arranged similar to those on dorsal surface. Tritubular pores lightly scattered over venter, except for head region. Body setae like those on dorsum, possibly fewer in number. Circulus absent. Legs moderately short and stout, with well-developed spines; claw slender, without denticle on plantar surface; digitules merely short, acute setae. In California specimens, tarsus shows indication of division. Antennae 6-segmented, quite short, stout, 2 apical segments bearing enlarged sensory setae. Eyes rather small, protuberant, pigmented. Notes. This species is very closely related to Rhizoecus distinctus (Hambleton), an eastern United States species, but differs in having the inner row of pores of the anal ring clouded, whereas, in the species last mentioned, these pores are clear. Ferris (1963) states there is a possibility that R. distinctus is a synonym of R. associatus. If they be united, R. associatus has page precedence over R. distinctus.

TAXONOMY

OF

SPECIES

Fig. 149. Rhizoecus associatus (Hambleton), collected on roots of Gramineae, at Santa Cruz, Santa Cruz County, California. (Illustration prepared from type material.)

373

374

TAXONOMY

OF

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California Records (Map 139) Santa Cruz Co.: Santa Cruz, II-9-64, on Gramineae (D. R. Miller and J. A. Froebe, U C D ) .

Map 139. Distribution in California of Rhizoecus associatus (Hambleton).

Mann San FwKiteC' San Matto Santa Cruz

Rhizoecus bicirculus McKenzie, new species Trinity Ground Mealybug (Figure 150) Type Locality and Host. Type and paratype adult females collected 6 miles northeast Weaverville, Stoney Creek on Trinity Lake, Trinity County, California, on roots of Ceanothus pumilus (Rhamnaceae), May 22, 1963, October 11, 1963, by R. F. Wilkey. These are the only known specimens of the species. Type Material. Holotype adult female (single specimen on slide), and paratypes of the same sex deposited in collection of California State Department of Agriculture, Sacramento. Additional paratype adult females in museum, University of California, Davis. North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. According to the collector, this mealybug was lightly covered with a fine, powdery, wax secretion. The species feeds on the rootlets of its host. There is usually a loose powdery wax in the soil around the roots where the insects are feeding. Recognition Characters. Adult females, mounted, 1.10 to 1.25 mm long, 0.40 to 0.50 mm wide; body form quite elongate oval. Dorsum without cerarii; anal lobes with but 2 or 3 slender setae, unsclerotized basal area. Dorsal ostioles not too conspicuous, though present.

Trilocular pores sparsely distributed over dorsum. Multilocular disk pores absent. Tritubular pores very few, restricted to a submarginal broken series from eighth abdominal segment forward to head, 2 or 3 present in median area of eighth segment, absent elsewhere on dorsum. Very few small, short tubular ducts of oral-collar type situated along submargin of last 2 or 3 abdominal segments, absent elsewhere on dorsum. Body setae small, slender, somewhat scattered over surface. Anal ring apical, with irregular pores, some shaded, others clear; its 6 setae slightly longer than diameter of ring itself. Ventral multilocular disk pores absent. Trilocular pores distributed much the same as on dorsum. Tritubular pores either absent or with only 1 or 2 on submargin of last abdominal segment. Small, short, tubular ducts, similar in size and shape to those on dorsum, infrequently found along margin of last 2 or 3 abdominal segments, absent elsewhere on venter. Body setae short and slender, distributed similar to those on dorsum except fewer in thoracic area. Two circuli present (hence the specific name bicirculus), these situated on third and fourth abdominal segments. Legs small, short, and stout, claw slender, without denticle on plantar surface; digitules slender, apically slightly knobbed. Eyes present, though very small and inconspicuous. Notes. This species appears closely related to Rhizoecus cacticans (Hambleton) but differs in possessing 2 circuli; fewer dorsal body setae especially on abdomen; and knobbed digitules on claw. On the other hand, R. cacticans has only 1 circulus; numerous body setae on abdomen; and setose digitules on claw. California Records (Map 140) Trinity Co.: 6 miles northeast Weaverville, Stoney Creek on Trinity Lake, X - l l - 6 3 , on Ceanothus pumilus (roots) (R. F. Wilkey, C D A ) .

M a p 140. Distribution in California of Rhizoecus bicirculus McKenzie.

TAXONOMY

OF

SPECIES

Fig. 150. Rhizoecus bicirculus McKenzie, new species, collected on roots of Ceanothus pumilus (Rhamnaceae), 6 miles northeast Weaverville, Trinity County, California.

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Rhizoecus bituberculatus McKenzie Bituberculate Soil Mealybug (Figure 151) Rhizoecus bituberculatus McKenzie, 1960. Hilgardia 2 9 ( 1 5 ) : 7 4 3 - 7 4 5 , illus. Type Locality and Host. 4 miles west of Newcastle, Placer County, California, in soil under Quercus wislizenii (Fagaceae). North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. Since specimens were recovered from soil subjected to Berlese funnel, no data is available on external appearance of this mealybug. The species occurs in the soil where it feeds on the roots of its host. Recognition Characters. Adult females, mounted, 0.90 to 1.10 mm long, 0.35 to 0.40 mm wide; body form quite elongate oval. Dorsum with cerarii entirely absent; the anal lobes with several slender setae of various lengths, these set in a quite distinct, but small, area of sclerotization. Dorsal ostioles well developed, their rims sclerotized. Dorsal surface beset with relatively small numbers of trilocular pores. Multilocular disk absent on dorsum except for a very few situated along posterior border of eighth abdominal segment. Bitubular pores scattered over dorsum, rather numerous, as many as 44 observed. Oral collar tubular ducts lightly scattered over dorsal surface. Body setae slender, of various lengths, arranged in rather widely separated areas particularly in thoracic region, on abdomen between segments, and in lateral patches. Anal ring borne at posterior apex of abdomen; its pores large, oval, and irregularly shaped, some appearing clear and open, others with varying amounts of pigmentation; its 6 setae but little longer than greatest diameter of ring. Ventral multilocular disk pores of 2 slightly different sizes, the larger ones present on apical to posterior border of seventh abdominal segment; smaller ones situated on abdominal segments 6 to 3, scattered in patches on thorax and head. Trilocular pores on abdomen and thorax confined to isolated patches. Bitubular pores few, similar in size to those on dorsum, confined to series along body margin. Oralcollar tubular ducts lightly scattered over ventral surfaces. Body setae short, slender and sparse. Normally with only 1 conical circulus, this situated on morphologically fourth abdominal segment (1 of 5 specimens in type series has 2 conical circuli evident, one on third and fourth segments, the one on fourth segment largest). Legs relatively short, stout, tibiae and tarsi armed with stout spines; claw with denticle on plantar surface; digitules setose, extending approximately to tip of claw. Antennae 6-segmented, quite short and stout, apical segment bearing 3 sensory setae, interantennal space equal to slightly more than width of basal segment. Eyes absent. Notes. This species is perhaps most closely related

to Rhizoecus spinosus McKenzie, but differs mainly in the possesion of ventral multilocular disk pores on thorax, whereas, in the species last mentioned, these structures are absent in this area. California Records (Map 141) El Dorado Co.: Near Nashville, 1-25-58, "in soil" (L. M. Smith and R. O. Schuster, U C D ) ; 3 miles north Placerville, V-18-61, "in soil" (R. O. Schuster, UCD). Placer Co.: 4 miles west of Newcastle, 111-21-58, "in soil" under Quercus wislizenii (W. H. Lange, L. M. Smith, and R. O. Schuster, U C D ) ; 1V-15-58, "in soil" (L. M. Smith and R. O. Schuster, U C D ) .

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Rhizoecus browni McKenzie Brown's Soil Mealybug (Figure 152) Rhizoecus browni McKenzie, 1961. Hilgardia 3 1 ( 2 ) : 43-45, illus. Type Locality and Host. 1 mile north Cajon Pass, San Bernardino County, California, in soil under Juniperus sp. (Cupressaceae). North American Distribution. Known only from California. Additional Hosts. A Berlese funnel was used to recover this mealybug from the soil. In addition to juniper, the soil from which this mealybug was taken supported 2 other plant species, namely, Sitanion hystrix (Gramineae) and Eriogonum jasciculatum ssp. polifolium (Polygonaceae). It is suspected that this Rhizoecus was feeding on the roots of the Sitanion grass, although additional field evidence is needed to substantiate this. Additional paratype specimens

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Fig. 151. Rhizoecus bituberculatus McKenzie, collected in soil under Quercus lizenii (Fagaceae), west of Newcastle, Placer County, California.

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Fig. 152. Rhizoecus browni McKenzie, collected "in soil" associated with grass (Gramineae), northwest of Spanish Flat, Napa County, California. (Illustration prepared from type material.)

TAXONOMY of this species were collected in soil under Quercus wislizenii (canopy), 5 miles northwest of Spanish Flat, Napa County, California. The soil supported 3 plant species: a grass, Stipa lepida (Gramineae), Claytonia lanceolata (Portulacaceae), and Dodecatheon hendersonii (Primulaceae). It is suspected that this mealybug was feeding on the roots of the grass, although additional field evidence is needed to substantiate this suspicion. External Features and Habitat. No information available other than that the mealybug occurs in the ground where it presumably feeds on the roots of its host. It is suspected that the grass roots of Sitanion hystrix and Stipa lepida may be the hosts attacked by this mealybug. Recognition Characters. Adult females, mounted, average approximately 1.00 mm long, 0.40 mm wide; body form quite elongate oval. Dorsum with cerarii entirely absent; the anal lobes with several slender setae of various lengths, these arising from slightly sclerotized area. Dorsal ostioles well developed. Trilocular pores evenly beset on dorsum. Multilocular disk pores present in small numbers along posterior margin of sixth and eighth abdominal segments. Tritubular pores numerous, scattered over dorsum, as many as 34 or 35 observed. A few small oral-collar type tubular ducts on certain abdominal segments. Body setae small, slender, evenly beset on dorsum. Anal ring apical; its pores large, oval and irregular shaped, clear and open; its 6 setae approximately twice as long as greatest diameter of ring. Ventral multilocular disk pores present in some numbers from apical to posterior margin of sixth abdominal segment, few situated near middle and front coxae, several located near mouthparts. Trilocular pores distributed very much as on dorsum, except on thorax confined to isolated patches showing as "clear" areas. Tritubular pores few on abdomen (7 actually counted on 1 side), very few along body margin, 1 or 2 situated near mouthparts, all as large as those on dorsum. Very few small oral-collar tubular type ducts scattered over venter. Body setae short, slender, intermixed with trilocular pores. Normally with 1 small, round, conical circulus present on fourth abdominal segment. Legs proportionately large and long for members of this genus; claw without denticle on plantar surface; digitules long and slender, apically swollen, as long as claw itself. Antennae 6-segmented, quite short, stout, apical segment bearing 4 sensory setae; interantennal space equal to approximately twice width of basal segment. Mouthparts short and broad; sclerotized patch anterior to clypeus. Eyes present. Notes. This mealybug resembles Rhizoecus cacticans (Hambleton), but differs in possessing ventral multilocular disk pores on abdomen, whereas, in the species last mentioned, these structures are absent anywhere on body. California Records (Map 142) Napa Co.: 5 miles northwest of Spanish Flat, III-

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1-61, "in soil" under Quercus wislizenii (H. L. McKenzie, U C D ) ; III-1-61, "in soil" associated with grass (R. O. Schuster and H. L. McKenzie, U C D ) . San Bernardino Co. : 1 mile north Cajon Pass, II5-60, "in soil" under Juniperus sp. (S. W. Brown, UCD).

Rhizoecus cacticans (Hambleton) Cacticans Mealybug (Figure 153; Color Plate IV) Ripersiella cacticans Hambleton, 1946. Rev. de Ent. 1 7 ( 1 - 2 ) : 6 4 - 6 6 , illus. Rhizoecus cacticans (Hambleton), Ferris, 1953. Atlas of scale insects of North America, Vol. VI:432-433, illus. Rhizoecus epiphyllis Ferris, McKenzie, 1960. Hilgardia 2 9 ( 1 5 ) : 7 4 5 and 747. Rhizoecus cacticans (Hambleton), McKenzie, 1960. Ibid. Rhizoecus cacticans (Hambleton), Williams, 1962. Bull. Brit. Mus. (Nat. Hist.) Ent. 12(1 ) : 4 3 . Type Locality and Host. Cayambe, Ecuador, on velvet grass, Holcus lanatus (Gramineae). North American Distribution. Recorded only from California. Additional Hosts. Bromus catharticus (Gramineae), Cyperus rotundus (Cyperaceae), Cyperus sp. (Cyperaceae), Distichlis spicata (Gramineae), Echeveria sp. (Crassulaceae), Epiphylum sp. (Cactaceae), Hamatocactus setispinus (Cactaceae), Kleinia sp. (Compositae), Lobivia shaferi (Cactaceae), Lobivia sp. (Cactaceae), Lolium perenne

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Fig. 153. Rhizoecus cacticans (Hambleton), collected on nut grass, Cyperns rotundus (Cyperaceae), at El Centro, Imperial County, California. (Illustration prepared from specimens collected on Lolium perenne [Gramineae], at Cayambe, Ecuador.)

TAXONOMY (Gramineae), Mammillaria sp. (Cactaceae), Phyllocactus sp. (Cactaceae), Saintpaulia ionantha (Gesneriaceae), and Sempervivum sp. (Crassulaceae). External Features and Habitat. According to Hambleton (1946), the nature of the waxy secretion of this mealybug is unknown. It is reported as viviparous. The species occurs on the roots (see col. pi. IV). Recognition Characters. Adult females, mounted, 1.20 to 2.60 mm long, 0.50 to 1.30 mm wide; body form elongate oval, rather stout. Dorsum without cerarii; anal lobes with but 3 slender setae, unsclerotized basal area. Dorsal ostioles very obscure, but present. Trilocular pores numerous over dorsum. Multilocular disk pores absent. Tritubular pores very small, not more than one and one-half diameter of trilocular pore, pore tubes strongly divergent apically. Very few exceedingly small, strongly sclerotized, tubular ducts mingled with trilocular pores. Body setae small, slender, evenly beset on dorsum. Anal ring comparatively large, with irregular pores; its 6 setae somewhat longer than greatest diameter of ring. Ventral multilocular disk pores absent. Trilocular pores distributed much the same as on dorsum. Tritubular pores few, confined to marginal areas. Small tubular ducts similar in size and shape to those on dorsum, likewise distributed. Body setae short and slender, intermixed with trilocular pores. Circulus very small, shaped like truncate cone when viewed in profile, situated on fourth abdominal segment. Legs well developed; claw slender; digitules elongate, weakly knobbed, extending to or beyond tip of claw itself. Antennae 6-segmented, quite short, stout, apical segment bearing several enlarged sensory setae. Eyes present. Notes. This species appears closely related to Rhizoecus browni McKenzie, but the absence of multilocular disk pores precludes confusion with that species. In R. browni multilocular disk pores are evident on both surfaces of last 3 or 4 abdominal segments; ventrally near mouthparts; and near fore- and mid-coxae. California Records (Map 143) Alameda Co.: Berkeley, XI^4~58, on Hamatocactus setispinus (H. L. McKenzie and A. E. Pritchard, UCD) ; VI-7-48, on Mammillaria sp. (C. A. Hanson, UCD); Niles, 11-25-48, on "cactus" (A. E. Pritchard, UCD). Imperial Co.: El Centro, 11-25-63, on Cyperus rotundus (G. O. Poinar and D. W. Ricker, CD A). Los Angeles Co.: West of Los Angeles, XI-30-61, on Echeveria sp. (A. G. C. Beresford, UCD). Riverside Co.: Norco, X-4-65, on Sempervivum sp. (N. W. Getz and J. H. Preston, CDA). Sacramento Co.: Sacramento (nursery), IX-22-65, on Echeveria sp. and Haworthia sp. (P. H. Siebert, CDA); IX-22-65, on Sempervivum sp. (S. Scribner, CDA). San Bernardino Co.: Ontario, X-26^+8, on Epiphylum sp. (C. R. Tower, CDA).

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San Diego Co.: Leucadia, II-9-55, on Kleinia sp. (D. Sprague and J. Wilson, CDA); 11-14-55, on Echeveria sp. (D. Sprague, CDA). Santa Cruz Co.: Santa Cruz, 111-26-64, on "succulents" (Rubra ray) (R. W. Nutter, CDA). Solano Co.: Near Grizzley Island, X-20-59, on Distichiis spicata (R. O. Schuster, UCD).

Rhizoecus californicus Ferris California Ground Mealybug (Figure 154) Rhizoecus californicus Ferris, 1953. Atlas of scale insects of North Amercia, Vol. VI: 434-43 5, illus. Type Locality and Host. Berkeley, Alameda County, California, collected "in soil under chaparral." North American Distribution. Known only from California. Additional Hosts. Most of the collections of this mealybug were made from soil samples placed in a Berlese funnel. Therefore, in most instances an actual host association has not been determined. A few records of actual hosts infested by this root mealybug are as follows: Artemisia douglasiana (Compositae), chaparral possibly referring to Ceanothus cuneatus (?) (Rhamnaceae), Dipsacus sp. (Dipsaceae), grass (Gramineae), and Polygonum sp. (Polygonaceae). External Features and Habitat. This mealybug is lightly dusted with a white, waxy secretion. The body color is white, approximately the same as the waxy secretion, and the legs are white and inconspicuous.

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Fig. 154. Rhizoecus californicus Ferris, collected "in soil under chaparral," at Berkeley, Alameda County, California. (Illustration prepared from type material.)

A seemingly smooth ovisac is formed which covers all but the anteriormost portion of the head. The mealybug infests the roots of its host. Recognition Characters. Adult females, mounted, 1.20 to 1.50 mm long, 0.75 to 1.00 mm wide; body form elongate oval, rather stout. Dorsum without cerarii; anal lobes bearing 3 slender, radiating setae, these set in a small, sclerotized area. Dorsal ostioles very distinct. Trilocular pores numerous, rather evenly beset on dorsum. Multilocular disk pores wanting. Tritubular pores relatively quite small and with tubes projecting but little from body wall, situated in transverse rows across abdominal and thoracic segments, scattered on head, the transverse rows on abdomen containing 8-10 pores, total number on dorsum approximately 100. Tubular ducts absent. Body setae short and slender. Anal ring apical; its 6 setae almost twice as long as diameter of ring. Ventral multilocular disk pores very few, confined to area about vulva forward to posterior margin of seventh segment, total pores approximately 30. Trilocular pores numerous, certain "clear" areas evident on thorax where these pores are absent. Tritubular pores, essentially of same size as those on dorsum, but fewer and scattered. Body setae short and slender, scattered on venter. Circulus very small, circular, when seen in profile appearing dome-shaped, slightly projected, this morphologically on fourth abdominal segment. Legs quite small; claw slender, without denticle; digitules exceeding apex of claw and slightly knobbed. Antennae 5-segmented. Very small sclerotized area situated anterior to clypeus. Eyes present. Notes. This species of Rhizoecus is readily distinguished from other species of this genus by its 5-segmented antennae. Aside from this character, it seems most closely related to Rhizoecus sonomae McKenzie, but differs in absence of dorsal multilocular disk pores on eighth abdominal segment as well as on venter of thorax, whereas, in the species last mentioned, these structures are present in these areas. California Records (Map 144) Alameda Co.: Berkeley, IV-23-46, "from soil" (H. M. Butterfield, CAS); III-2-58, "in soil under chaparral" (L. M. Smith, UCD). Marin Co.: 2 miles northeast Bolinas, VIII-27-63, on Artemisia douglasiana (roots) (D. R. Miller, UCD); 1 mile west Bolinas, 1-19-60, "in soil" (W. G. litis and C. L. Judson, UCD) ; 6 miles east Point Reyes, III—1—60, on Dipsacus sp. (L. M. Smith, UCD); III—1—60, "in soil" (R. O. Schuster, UCD). San Mateo Co. : Millbrae, San Andreas Lake, X I 1957, "in soil in grassland" (D. W. Price, UCD). Yolo Co.: Davis, IX-23-61, on Polygonum sp. (M. E. Irwin, UCD).

Rhizoecus cyperalis (Hambleton) Viviparous Ground Mealybug (Figure 155) Morrisonella cyperalis Hambleton, 1946. Rev. de Ent. 17(1-2):22-23, illus. Coccidella cyperalis (Hambleton), Hambleton, 1946. Proc. Biol. Soc. Wash. 59:177. Rhizoecus cyperalis (Hambleton), Ferris, 1953. Atlas of scale insects of North America, Vol. VI:438-439, illus. Type Locality and Host. Santiago de Maria, El Salvador, on small sedge, Cyperus tenerrimus (Cyperaceae). North American Distribution. California and El Salvador. Additional Hosts. Adiantum sp. (Polypodiaceae). External Features and Habitat. According to Hambleton (1946), who collected this species, the adult females are lightly covered with a powdery wax and are viviparous. Like other members of this genus, it occurs on the roots of its host. Recognition Characters. Adult females, mounted, 1.42 to 1.90 mm long, 1.10 to 1.58 mm wide; body form elongate oval, rather stout. Dorsum without cerarii; anal lobes with 4 or 5 slender setae of various lengths, unsclerotized basal area. Dorsal ostioles well developed, their rims sclerotized. Trilocular pores rather evenly beset on dorsum. Multilocular disk pores absent. Tritubular pores with each loculus as large as or larger than trilocular pore, rather few, total number scarcely exceeding 25, scattered over dorsal surface. Tubular ducts absent. Body setae small, evenly beset on dorsum. Anal ring apical; pores of inner and outer rings rather narrow, elon-

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& Fig. 155. Rhizoecus cyperalis (Hambleton), collected on a fern, Adiantum sp. (Polypodiaceae), at Colma, San Mateo County, California. (Illustration prepared from type material.)

TAXONOMY gate, clouded; its 6 setae little longer than diameter of ring itself. Ventral multilocular disk pores very few, there being scarcely more than 6, these concentrated around vulva. Trilocular pores distributed much as on dorsum. Tritubular pores very few, confined to a series along body margins, these as large as those on dorsum. Tubular ducts absent. Body setae short and slender. Circulus absent. Legs quite small; claw slender, without denticle or tooth on plantar surface; digitules very short, apically acute. Antennae 6-segmented, very widely spaced near apex of head. Eyes present. Notes. This species is closely related to Rhizoecus nemoralis (Hambleton), but is distinguished from it by possessing 2 elongate setae and 2 or 3 smaller auxiliary setae in anal lobe area, and 3-4 body setae usually surrounding each tritubular pore. R. nemoralis, on the other hand, has 3 elongate setae and 5-8 smaller auxiliary setae in anal lobe area, and the body setae are not necessarily associated with the tritubular pores. California Records (Map 145) San Mateo Co.: Colma, 11—24—55, on Adiantum sp. (W. E. Davis, CD A).

Rhizoecus disjunctus McKenzie, new species Unipored Mealybug (Figure 156) Type Locality and Host. Type and paratype adult females collected at Corona (Prado Dam), Riverside County, California, on Eriogonum sp. (Polygona-

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ceae), December 27, 1964, by D. R. and J. F. Miller. These are the only known specimens of the species. Type Material. Holotype adult female (single specimen on slide) and female paratypes in museum, University of California, Davis. Additional paratypes in United States National Collection of Coccoidea (U. S. National Museum), Washington, D. C., and in collection of California State Department of Agriculture, Sacramento. North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. This mealybug is lightly dusted with a smooth, white, waxy secretion. The body form is elongate oval; body color whitish; legs white. There are no visible filaments. No ovisac is produced. The mealybug feeds on the roots of its host. There were no ants found in association with the mealybug. Recognition Characters. Adult females, quite small for members of this genus, mounted, 0.75 to 1.10 mm long, 0.40 to 0.45 mm wide; body form elongate oval. Dorsum without cerarii; anal lobes with but 2 or 3 slender setae, unsclerotized basal area. Dorsal ostioles present. Trilocular pores rather evenly beset on dorsum. A few multilocular disk pores present along posterior margin of seventh and eighth abdominal segments, absent elsewhere on dorsal surface. Unitubular pores small, heavily sclerotized, some appearing conicalshaped, others with slight bifurcate tip (see figure 156), with tube projecting from body derm, occurring in transverse rows across abdominal and thoracic segments, scattered on head. Oral-collar type tubular ducts short, with rather heavily sclerotized orifices, scattered over dorsal surface. Body setae short, slender, fairly evenly distributed on dorsum. Anal ring comparatively large, with irregular cells, some pigmented, others clear; its 6 setae somewhat longer than greatest diameter of ring. Ventral multilocular disk pores very few, these situated on apical segment of abdomen forward to posterior margin of sixth segment. Trilocular pores distributed much the same as on dorsum, except for certain "clear" areas on thorax and submargin of abdomen. Unitubular pores similar in size and shape to those on dorsum somewhat scattered on last 3 abdominal segments, and in submarginal irregular band from sixth abdominal segment forward to head. Body setae short and slender, intermixed with trilocular pores. Circulus small, shaped like a truncate cone when viewed in profile, situated on fourth abdominal segment. Legs well developed, stout; claw slender, without denticle on plantar surface; digitules slender, apically slightly knobbed, as long as claw itself. Antennae 6-segmented, quite short though stout, sensory setae borne on apical 2 segments. Eyes present. Notes. This species shares with Rhizoecus geniculates James, an east African species, the possession

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Fig. 156. Rhizoecus disjunctus McKenzie, new species, collected on rootlets of Eriogonum sp. (Polygonaceae), at Corona (Prado Dam), Riverside County, California.

TAXONOMY of the peculiar type of unitubular pores, but differs from the species last mentioned in having only a few multilocular disk pores confined to last 3 or 4 abdominal segments on both surfaces of body. Rhizoecus geniculatus, on the other hand, has numerous multilocular disk pores widely scattered on both sides of body. The presence of the unitubular pores in Rhizoecus disjunctus make this species somewhat atypical for members of this genus. However, all other characteristics are quite normal for Rhizoecus, and it is, therefore, considered to be a genuine component of this group. California Records (Map 146) Riverside Co.: Corona (Prado Dam), XII-27-64, on Eriogonum sp. (rootlets) (D. R. Miller, UCD).

Rhizoecus eluminatus McKenzie Blind Mealybug (Figure 157) Rhizoecus eluminatus McKenzie, 1960. Hilgardia 29(15): 747-749, illus. Type Locality and Host. Bayside, Humboldt County, California, on Sinningia sp. ( = Gloxinia sp.) (Gesneriaceae). North American Distribution. Known only from California. Additional Hosts. Achillea ageratifolia (Compositae). External Features and Habitat. No available information on appearance of this species in life.

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Specimens of the mealybug were collected from the roots of its host. Recognition Characters. Adult females, mounted, 1.60 to 2.00 mm long, 0.70 to 1.40 mm wide; body form elongate oval, rather stout. Dorsal ostioles very distinct, their rims sclerotized. Dorsum with cerarii entirely absent, anal lobes with 4 or 5 slender setae of various lengths, no sclerotization. Trilocular pores rather numerous, evenly distributed over dorsum. Multilocular disk pores absent. Tritubular pores scattered over dorsum, rather few, total number scarcely exceeding 28. Tubular ducts absent. Body setae small and slender. Anal ring borne close to posterior apex of abdomen; coarsely cellular; its 6 setae slightly longer than greatest diameter of ring. Ventral multilocular disk pores very few, situated in area of vulva forward to posterior boarder of seventh abdominal segment, observed range from 13 to 17, average 14. Trilocular pores possibly less numerous than on dorsum, evenly distributed. Tritubular pores few, apparently confined to a series along body margins, these as large as those on dorsum. Tubular ducts absent. Ventral body setae short and slender. Circulus absent. Legs relatively short; claw slender, without denticle on plantar surface; digitules very short, apically acute. Antennae 6-segmented, quite stout, their bases rather widely spaced for members of this genus, separated by more than twice width of basal segment. Eyes absent. Notes. This species is most closely related to Rhizoecus pritchardi McKenzie, but differs from it principally in the complete absence of eyes, whereas in the species last mentioned eyes are definitely present. Beyond this, when specimens of these species are placed side by side under 1 ocular, the antennae of P. pritchardi are defintely more slender than those of P. eluminatus, and the beak is just a little shorter in the first, with its tip rounded, in contrast to the very slightly longer beak with bluntly angulate tip in the second. The legs seem much alike, except the femora are recognizably not quite so stout in the first; the anal lobes and the anal ring setae appear practically identical; the number of multiloculars in the ventral abdominal region seem to be about the same; and the distribution pattern of the titubular pores seems much the same, but with enough variability to create uncertainty as to the true average pattern. The presence or absence of eyes in these 2 species seems most significant and provides an excellent key character separating them. California Records (Map 147) Alameda Co.: Berkeley (University of California Botanical Gardens), V-1959, on Achillea ageratifolia (A. E. Pritchard, UCD). Humboldt Co.: Bayside, IX-24-54, on Sinningia (= Gloxinia) sp. (W. D. Thomas, CDA).

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Fig. 157. Rhizoecus eluminatus McKenzie, collected on gloxinia, Sinningia sp. ( = Gloxinia sp.) (Gesneriaceae), at Bayside, Humboldt County, California.

TAXONOMY

Rhizoecus falcifer Kiinckel d'Herculais Ground Mealybug (Figure 158; Color Plate X X I ) Rhizoecus falcifer Kiinckel d'Herculais, 1878. Ann. Ent. Soc. de France (ser. 5) 8:161-164, illus. Ripersia terrestris Newstead, 1895. Ent. Monthly Mag. 31:213, 236, illus. Rhizoecus (?) terrestris (Newstead), Fernald, 1903. Mass. Agr. Expt. Sta. Spec. Bui. 88, p. 114. Rhizoecus africanus Brain, 1915. Trans. Roy. Soc. So. Africa 5 ( 2 ) : 2 5 . Rhizoecus decoratus Green, 1926. Ent. ' Monthly Mag., 62:177, illus. Rhizoecus moruliferus Green, 1933. Stylops 2 ( 3 ) : 52-53, illus. Rhizoecus falcifer Kiinckel d'Herculais, Rau, 1937. Jour. N.Y. Ent. Soc. 4 5 ( 2 ) : 2 6 7 . Rhizoecus falcifer Kiinckel d'Herculais, Hambleton, 1946. Rev. de Ent. 17(1-2) :53-56, illus. Rhizoecus falcifer Kiinckel d'Herculais, Ferris, 1953. Atlas of scale insects of North America, Vol. VI: 444-445, illus. Rhizoecus moruliferus Green, Williams, 1961. Ent. Monthly Mag. 97:93. Rhizoecus falcifer Kiinckel d'Herculais, Williams, 1962. Bull. Brit. Mus. (Nat. Hist.) Ent. 12(1 ) :47. Type Locality and Host. Paris, France on a palm, Seaforthia elegans (= Ptychosperma elegans) (Palmaceae). North American Distribution. California, Missouri, New Jersey, and New York. Additional Hosts. This mealybug occurs on a wide range of hosts a few of which are recorded as follows:

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Abutilon sp. (Malvaceae), Acacia sp. (Leguminosae), Adiantum sp. (Polypodiaceae), Agrostis sp. (Gramineae), Aquilegia sp. (Ranunculaceae), Anemone sp. (Ranunculaceae), Buxus sempervirens and Buxus sp. (Buxaceae), Carex sp. (Cyperaceae), Chrysanthemum frutescens and Chrysanthemum spp. (Compositae), Cistus sp. (Cistaceae), Citrus sinensis (Rutaceae), Codiaeum sp. (Euphorbiaceae), Coffea liberica (Rubiaceae), Convolvulus arvensis (Convolvulaceae), Cynodon dactylon (Gramineae), Cyperus sp. (Cyperaceae), Delphinium sp. (Ranunculceae), Dracaena sp. (Agavaceae), Erodium sp. (Geraniaceae), Fragaria sp. (Rosaceae), Gardenia intermedia (Rubiaceae), grasses (various) (Gramineae), Howea belmoreana and Howea sp. (Palmaceae), Jasminum sp. (Oleaceae), Kentia sp. (Palmaceae), Ligustrum sp. (Oleaceae), Lotus sp. (Leguminosae), "moss lawn," Ophiopogon sp. (Liliaceae), palms (various) (Palmaceae), Pelargonium sp. (Geraniaceae), Petunia sp. (Solanaceae); Phoenix robelenii (Palmaceae), Phormium sp. (Agavaceae), Picea sp. (Pinaceae), Piper sp. (Piperaceae), Prunus persica and Prunus spp. (Rosaceae), Ribes sp. (Saxifragaceae), Sabal umbraculifera (= Blackburnia) (Palmaceae), Senecio (= Kleinia) pendula (Compositae), Stephanotis sp. (Asclepiadaceae), Syringa vulgaris (Oleaceae), Thymus sp. (Labiatae), Tropaeolum majus and Tropaeolum spp. ( = Nasturtium of gardeners) (Tropaeolaceae), Vitis sp. (Vitaceae), Watsonia sp. (Iridaceae), and weeds. External Features and Habitat. The species secretes a small amount of wax which, when the insects are numerous, gives the soil a somewhat bluish appearance. In life this mealybug resembles some of the larger forms of Collembola. It occurs on the roots of its host and in the soil about the roots (see col. pi. XXI). Recognition Characters. Adult females, mounted, 2.40 to 3.90 mm long, 1.00 to 1.60 mm wide; body form quite elongate oval. Dorsal ostioles very distinct, bordered by sclerotized rims. Dorsum without cerarii; anal lobes distinctly projecting, bearing 5 - 7 long and radiating setae, 3 of them much longer and stouter than others, unsclerotized. Trilocular pores thickly beset over entire dorsum. Multilocular disk pores very few on dorsum, these confined mainly to anterior half of body. Tritubular pores large, numerous, with tubes very short, projecting but slightly from body, these pores appearing merely as extra large, strongly sclerotized, trilocular pores—total of more than 100 on dorsum alone. Tubular ducts absent. Body setae small, slender, thickly beset on dorsal surface. Anal ring apical, with few quite large, irregular pores; its 6 setae, slender, about twice as long as diameter of ring. Ventral multilocular disk pores extremely numerous especially about region of vulva, in 2 transverse rows across posterior margin of segment 7, and in more or less single rows along posterior margin of abdominal segments 5-3. Trilocular pores numerous, distributed similar to those on dorsum, certain "clear"

390

TAXONOMY

OF

SPECIES

Fig. 158. Rhizoecus falcifer Kiinckel d'Herculais, collected on iris, Iris sp. (Iridaceae), at Berkeley, Alameda County, California. (Illustration prepared from specimens collected on "lawn moss," at Oakland, Alameda County, California.)

COLOR PLATE XXI. The ground mealybug, Rhizoecus falcifer Kunckel d'Herculais, in soil on gladiolus bulbs (corms), Gladiolus sp. (Iridaceae), Sacramento, Sacramento County, California.

TAXONOMY

areas showing on thorax. Tritubular pores occur for most part in transverse rows on median area of abdominal segments 5-7, these pores slightly smaller than those on dorsum. Tubular ducts absent. Ventral body setae short and slender, numerous, distributed in same general pattern as trilocular pores. Circulus absent. Legs relatively large; claw slender, without denticle on plantar surface; digitules short, setose. Antennae 5-segmented. Small sclerotized area situated anterior to clypeus. Eyes apparently absent. Notes. The presence of numerous, large-sized, tritubular pores, especially on the dorsum, a 5-segmented antenna, and the absence of a circulus readily distinguishes this species from all other California Rhizoecus. This mealybug is one of the more common species of this group found in California. At times it becomes sufficiently abundant to cause injury to its host. California Records (Map 148)

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Alameda Co.: Berkeley, VII-8-44, on Iris sp. (E. O. Essig, USNM); IX-25-59, on Chrysanthemum sp. (L. A. Falcon, U C D ) ; 11-15-59, on Pelargonium sp. (A. E. Pritchard, U C D ) ; Oakland, 1-29-38, on Agrostis sp. (E. O. Essig, U C D ) ; VII-19-39, in "moss lawn" (E. O. Essig, CAS); IV-30-52, on Heuchera sp. (G. B. Laing, U C D ) ; Piedmont, V-10-59, on ? (K. S. Hagen, U C D ) . Contra Costa Co. : El Cerrito, 11-23-60, "in soil" (L. Wiegmann, U C D ) . Kern Co.: Bakersfield, XII-19-34, on Kentia sp. (C. S. Morley, C D A ) . Los Angeles Co.: Catalina Island, VI-21-29, on

OF

SPECIES

391

Kentia sp. (roots) (K. L. Wolff, L E M ) ; Gardena, V-26-58, on Ophiopogon sp. (Hever, U C D ) ; Inglewood, II-4-44, on Hibiscus sp. (soil) (Lichty, LEM); Los Angeles, VI-9-17, on Chrysanthemum sp. (R. S. Woglum, U C D ) ; VI1I-17-25, on Tropaeolum (= Nasturtium) sp. (L. E. Myers, U C D ) ; X I I 10-28, on "fern" (roots) (L. E. Myers, U C D ) ; Long Beach, IV-1941, "in soil" (collector ?, L E M ) ; Montebello, IV-16-31, on Phoenix robelenii (L. S. Jones, U C D ) ; Montebello (lath house), 1-17-35, on Phoenix canariensis (roots) (collector ?, U C D ) ; Whittier, VII-29-33, on Anthemis tinctoria (F. S. Stickney, USNM). Marin Co.: Fairfax, V - 5 - 5 8 , "in soil" (W. Erickson, U C D ) ; San Anselmo, VII-20-59, in soil under Picea sp. (J. L. Joos, U C D ) ; San Rafael, V I I I 1959, in soil (R. S. Hunsinger, UCD). Monterey Co.: Salinas, IV-17-62, "in soil" (L. Lanini, UCD) ; V-19-62, on "hedge"? (W. H. Lange and L. Lanini, U C D ) . Placer Co.: Colfax, V-10-56, "in soil" (M. D. Norman, C D A ) . Sacramento Co.: Sacramento, X - 2 6 - 4 2 , on Anemone japonica (H. H. Keifer, C D A ) ; II-7-40, on Buxus sempervirens (J. B. Steinweden, C D A ) ; I I 17-41, on Kentia sp. (roots) (J. B. Steinweden, C D A ) ; IV-27-63, on Watsonia sp. (H. L. McKenzie, U C D ) ; Walnut Grove, XI-10-59, "in soil" (Rogers, U C D ) . San Diego Co.: La Jolla, V-18-63, "in soil" (R. A. Lewin, U C D ) ; La Mesa, IV-5-57, on Matthiola sp. (K. Sims and D. Brown, C D A ) ; National City, 11-23-55, under "flower pot" (G. G. Beevor, C D A ) ; Pacific Beach, VIII-30-56, on Jasminum sp. (J. H. Wilson, C D A ) ; VII-29-53, under "flower pots" (G. Schwegel, C D A ) ; San Diego, V - 7 - 6 4 , on "palm" Aralia sp. (soil) (J. R. Carlin, C D A ) . San Francisco Co.: San Francisco, X - 3 1 - 6 0 , on Escallonia rubra (roots) (M. Stufflebeam, C D A ) . San Joaquin Co.: Stockton, X - 6 - 5 3 , on ? (roots) (D. Zuckswert, C D A ) . San Mateo Co.: San Carlos, VIII-4-47, on Fragaria sp. (roots) (C. M. Sill, C D A ) . Santa Barbara Co.: Santa Barbara, III—31—42, on Lotus sp. (J. B. Steinweden and S. Smith, U C D ) ; VII-22-43, on ? (roots) (C. E. Ehmann and R. P. Allen, C D A ) . Santa Clara Co.: Palo Alto, 111-25-41, "in soil" (Duncan, UCD); San Jose, 1-11-21, Chrysanthemum frutescens (L. R. Cody, C A S ) ; 1-11-21, on Buxus sp. (E. O. Essig, CAS); IV-13-21, on Buxus sp. (R. D. Hartman, USNM); IX-25-59, on Fragaria sp. (W. W. Allen, U C D ) . Santa Cruz Co.: Santa Cruz, 11-27-33, on ? (roots) (C. V. Dick, C D A ) ; Watsonville, IV-19-57, on Agapanthus sp. (L. R. Gillogly, C D A ) . Solano Co.: Vallejo, VII-13-44, on "Calla Lily" (E. O. Essig, U S N M ) ; V - 2 7 - 5 5 , "in soil" (L. A. Black, CDA) ; Rio Vista, IX-14-64, on Gardenia sp. (roots and soil) (W. H. Lange, C D A ) . Sonoma Co.: Healdsburg, II-1-62, "in soil" (W.

392

TAXONOMY

OF

SPECIES

R. Michie, UCD); Santa Rosa, V-15-61, in soil (M. E. Imsdale, UCD). Stanislaus Co.: Modesto, X-5-56, on Ajuga sp. (G. E. White, CDA). Ventura Co.: Oxnard, VI-1925, in soil (H. S. Smith, UCD); Santa Paula, X-14-33, in soil under Cupressus sp. (E. L. Smith, CDA); Ventura, XII15-33, on Veronica sp. (roots) (E. L. Smith, LEM). Yolo Co.: Davis, VIII-6-63, on Cynodon dactylon (W. H. Lange, UCD); V-24-62; on Aquilegia sp. and Jasminum sp. (F. E. Strong, UCD).

Rhizoecus gracilis McKenzie Slender Ground Mealybug (Figure 159) Rhizoecus gracilis McKenzie, 1961. Hilgardia 31(2) :45-47, illus. Type Locality and Host. Cahill Ridge, San Mateo County, California, in soil beneath chaparral. North American Distribution. Known only from California. Additional Hosts. Achillea millefolium (Compositae), Artemisia tridentata (Compositae), Atriplex sp. (Chenopodiaceae), and Chenopodiaceae. Mealybug also found in several soil collections, although actual host association undetermined. External Features and Habitat. No information other than that the mealybug inhabits the soil where it presumably assumes the role of a root feeder. Recognition Characters. Adult females, mounted, 0.95 to 1.00 mm long, 0.30 to 0.35 mm wide; body form quite elongate oval. Dorsal ostioles well developed. Dorsum with cerarii entirely absent. Anal lobe with several slender setae of various lengths, these arising from a slightly sclerotized area. Trilocular pores sparsely beset on dorsum. Multilocular disk pores in small numbers on eighth abdominal segment. Bitubular pores relatively few 12 to 16 scattered over abdomen, very few present along body margin on thorax and head. Several very small, short, oral-collar tubular ducts scattered mostly on abdomen, very few on thorax and head. Body setae small, sparsely beset on dorsal surface. Anal ring borne at posterior apex of abdomen; its pores quite large, elongate oval, irregularly shaped, clear and open; its 6 setae twice as long as greatest diameter of ring. Ventral multilocular disk pores in moderate numbers from posterior apex of body forward to seventh, infrequently sixth, abdominal segments, absent on thorax and head. Trilocular pores sparsely distributed on venter, confined to isolated patches, along with short body setae, showing as rather marked "clear" areas, particularly on thorax, less so on abdomen. One or two bitubular pores on abdomen, absent elsewhere ventrally. Very few small oral-collar tubular ducts scattered over venter. Body setae short, slender, sparsely distributed. Circulus small, round, with reticulated center, cone-shaped in profile (one specimen has 2 circuli,

anterior one smallest). Legs comparatively stout; claw without denticle on plantar surface; digitules long, slender, apically swollen, as long as claw itself. Mouthparts short, moderately broad. Antennae 6-segmented, quite short, stout, apical segment bearing 4 sensory setae; interantennal space equal to slightly less than width of basal segment. Sclerotized patch anterior to mouthparts. Eyes present. Notes. This species seems only remotely related to Rhizoecus solani (Hambleton) and R. totonicapanus (Hambleton). It differs from these species principally in the character of the bitubular pores which appear as small sclerotized cones, from the apex of which arise 2 curving tubes. In R. solani the bitubular pores appear in pairs, these consisting of a pair of tubular pores, the openings of which are set close together, the tubes parallel or somewhat divergent; in R. totonicapanus these structures are evident as strongly divaricating bitubular pores. California Records (Map 149)

Alpine Co.: Monitor Pass, VIII-9-64, "under rock" (D. R. Miller, UCD). Inyo Co.: 2 miles west of Independence, V-6-60, "in soil" (A. S. Menke and F. D. Parker, UCD). Lassen Co.: Susanville, V-17-58, "in soil" (R. W. Gerhardt, UCD). Orange Co.: Trabuco Canyon, 111-28-64, "in soil" (D. R. Miller and J. A. Froebe, UCD). San Bernardino Co.: 15 miles north Kramer Junction, XII-28-64, on Atriplex sp. (roots) (D. R. and J. L. Miller, UCD).

TAXONOMY

OF

SPECIES

Fig. 159. Rhizoecus gracilis McKenzie, collected in soil beneath chaparral, at Cahill Ridge, San Mateo County, California.

393

394

TAXONOMY

OF

SPECIES

San Diego Co.: Borrega Springs, 1-27-65, on Chenopodiaceae (roots) (D. R. Miller, UCD); 2 miles south Valley Center, 111-27-64, "in soil" (D. R. Miller and J. A. Froebe, UCD); Warner Springs, 1-26-65, on Artemisia tridentata (above ground under bark) (D. R. Miller, UCD). San Mateo Co.: Cahill Ridge, V-7-60, "in soil beneath chaparral" (D. W. Price, UCD); 10 miles north Pescadero, II-8-64, on Achillea millefolium (roots) (D. R. Miller and J. A. Froebe, UCD).

Rhizoecus graminis (Hambleton) Graminis Ground Mealybug (Figure 160) Morrisonella graminis Hambleton, 1946. Rev. de Ent. 17(1-2):28-29, illus. Coccidella graminis Hambleton, 1946. Proc. Biol. Soc. Wash. 59:177. Rhizoecus graminis (Hambleton), Ferris, 1953. Atlas of scale insects of North America, Ser. V:446-447, illus. Type Locality and Host. Oakland, Alameda County, California, on bent grass, Agrostis sp. (Gramineae). North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. No available information on external appearance of this mealybug. The species occurs on the roots of its host. Recognition Characters. Adult females, mounted, 1.26 to 1.74 mm long, 0.50 to 0.72 mm wide; body form elongate, oval. Dorsum without cerarii; anal lobes bearing but 2 slender setae, these set in a faint area of sclerotization. Dorsal ostioles well developed, their rims heavily sclerotized. Trilocular pores quite numerous, uniformly distributed on dorsum. Multilocular disk pores rather numerous, occurring in bands across the segments from eighth abdominal segment forward to head. Tritubular pores sparsely distributed over dorsum, total scarcely attaining 20 in number, each pore with loculus slightly larger than trilocular pore. Tubular ducts absent. Body setae variable in size, uniformly distributed. Anal ring apical, presenting no distinctive specific characters; each of its 6 setae almost twice as long as diameter of ring. Ventral multilocular disk pores fairly numerous, especially in bands across abdominal segments, scattered on thorax and head. Trilocular pores situated on abdomen much as those on dorsum, certain "clear" areas evident on thorax where these pores are absent. Tritubular pores few, confined for most part along margins of body. Body setae variable in size, uniformly distributed. Circulus absent. Legs well developed, moderately elongate; claw slender, moderately long curved, without denticle; digitules short, acutely pointed, barely attaining one-fourth length of claw. Antennae 6-segmented, rather slender, interantennal space equal to

approximately length of apical antennal segment. Small area of sclerotization anterior to clypeus. Eyes present. Notes. This mealybug is rather closely related to Rhizoecus neomexicanus McKenzie, a Mexican species, but differs from it mainly in possessing far fewer multilocular disk pores on both body surfaces. California Records (Map 150) Alameda Co.: Oakland, 1-29-38, on Agrostis sp. (E. O. Essig, CAS). San Mateo Co.: Colma, XII-12-63, "in soil" (C. S. Koehler, CDA).

Rhizoecus kondonis Kuwana Kondo Mealybug (Figure 161; Color Plate III) Rhizoecus kondonis Kuwana, 1923. (Japan) Dept. Agr. and Com., Imp. Plant Quar. Sta. Bui. 3:55-57, illus. Rhizoecus kondonis Kuwana, Hambleton, 1946. Rev. deEnt. 17(1-2) :56-57, illus. Rhizoecus kondonis Kuwana, Ferris, 1953. Atlas of scale insects of North America, Vol. VI:448449, illus. Rhizoecus kondonis Kuwana, McKenzie, 1960. Hilgardia 29(15) :749. Type Locality and Host. Wakayama, Japan, on orange, Citrus sp. (Rutaceae). North American Distribution. Known only from California. Additional Hosts. This species has been found on many hosts, a few of which include: Carex sp. (?)

TAXONOMY

OF

SPECIES

Fig. 160. Rhizoecus graminis (Hambleton), collected on bent grass, Agrostis sp. (Gramineae), at Oakland, Alameda County, California.

395

396

TAXONOMY

OF

SPECIES

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Spilococcus atriplicis (Cockerell) Southern California Mealybug (Figure 169) Dactylopius solani var. atriplicis Cockerell, 1895. Amer. Nat. 29:729-730. Pseudococcus solani var. atriplicis (Cockerell), Fernaid, 1903. Mass Agr. Expt. Sta. Spec. Bull. 88, p. 110. Spilococcus atriplicis (Cockerell), Ferris, 1953. Atlas of scale of insects of North America, Vol. VI:466, illus.

TAXONOMY

OF

SPECIES

Fig. 169. Spilococcus atriplicis (Cockerell), collected on arrow-weed, Pluchea sericea (Compositae), at Seeley, Imperial County, California. (Illustration prepared from type material.)

415

416

TAXONOMY

OF

SPECIES

Spilococcus eriogoni (Ehrhorn), Ferris, 1953. Ibid. (as a misidentification). Pseudococcus yerbae-santae Essig, Ferris, 1953. Ibid. (as a misidentification ). Spilococcus atriplicis (Cockerell), McKenzie, 1960. Hilgardia 29(15):761 (discussion under Spilococcus eriogoni [Ehrhorn]). Type Locality and Host. Mesilla (near Agricultural College), Dona Ana County, New Mexico, on shade scale, A triplex canescens (Chenopodiaceae). North American Distribution. California and New Mexico. Additional Hosts. Baccharis sp. (Compositae), Eriogonum inflatum (Polygonaceae), Eriogonum sp. (Polygonaceae), Haplopappus sp. (Compositae), andPluchea sericea (Compositae). External Features and Habitat. According to the original description, this species is "pale greenish, sparsely mealy, no lateral processes; forming no ovisac, but a cushion of white cottony matter in which are seen lively young." The male is winged. The mealybug occurs on the twigs, branches, and roots of its host. Recognition Characters. Adult females, mounted, 2.00 to 2.50 mm long, 1.30 to 1.60 mm wide; body form broadly oval. Dorsum with normally at least 15 pairs of recognizable cerarii, and possibly some specimens may show even 1 or 2 more; anal lobe cerarius with 2 moderately large, conical setae, 2 or 3 slender auxiliary setae of various lengths, loose cluster trilocular pores. Certain frontal cerarii may contain 3 or possibly 4 conical setae, while the others have but 2, these all quite small, no auxiliary setae, very slight concentration trilocular pores. Trilocular pores evenly beset on dorsum. Very few oral-collar tubular ducts evident on ninth abdominal segment. Oral-rim tubular ducts fairly numerous, arranged in a single row of as many as 12-14 across each abdominal segment, except the last, scattered over thoracic segments and head. Body setae short, slender, comparatively sparse. Anal ring with no unusual features; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores present from apical to posterior margin of fifth segment in midregion of abdomen. Trilocular pores about as numerous as those on dorsum, except in thoracic area. Oralcollar tubular ducts occupy same general area on abdomen as multilocular disk pores, rather numerous in lateral areas of eighth to sixth segments. Oral-rim tubular ducts of same size as those on dorsum, occur in small numbers in lateral areas of all segments from eighth abdominal segment forward to head, a few in sternal areas, and about spiracles. Body setae slender, comparatively sparse, some noticeably longer than those on dorsum. Circulus absent. Legs well developed; hind coxae with translucent pores; claw with or without small denticle on plantar surface. Antennae normally 8segmented. Notes. This species is very closely related to Spilo-

coccus eriogoni (Ehrhorn), and Ferris (1963) considered the 2 species to be identical. It is the author's (1960) opinion, however, that they are definitely distinct. This statement is based on the fact that S. atriplicis normally has at least 15 recognizable pairs of cerarii present, whereas in S. eriogoni the number of cerarii are much reduced, there being only 7 or 8 pairs counting forward from the anal lobes, with at times a recognizable ocular or frontal pair. In addition, S. eriogoni normally possesses almost twice as many dorsal oral-rim tubular ducts, particularly on the fifth and sixth abdominal segments, as compared to S. atriplicis. In view of this development, the author believes Ferris' opinion as expressed in his 1953 key to species, where he actually separated S. atriplicis and S. eriogoni, is sounder than his statement under the discussion of S. atriplicis that the 2 are identical. California Records (Map 159) Imperial Co.: Brawley, VII-8-64, on Baccharis sp. (J. Taylor and C. Gammon, CDA); Seeley, 1-24-57, on Pluchea sericea (G. G. Beevor, CDA); Thermal, XII-6-56, on Eriogonum inflatum (R. P. Allen, CDA). San Bernardino Co.: 9 miles south Adelanto, VII23-63, on Eriogonum sp. (foliage) (D. R. Miller, UCD).

Spilococcus cactearum McKenzie Cactus Mealybug (Figure 170; Color Plate II) Spilococcus cactearum McKenzie, 1960. Hilgardia 29(15):756-761, illus. Spilococcus cactearum McKenzie, 1961. Ibid. 31 ( 2 ) : 47.

TAXONOMY

OF

SPECIES

Fig. 170. Spilococcus cactearum McKenzie, collected on horse crippler cactus, Homalocephala texensis (Cactaceae), at Berkeley (University of California Botanical Gardens), Alameda County, California.

417

418

TAXONOMY

OF

SPECIES

Spilococcus cactearum McKenzie, Williams, 1962. Bull. Brit. Mus. (Nat. Hist.) Ent. 12(1):5254, illus. Type Locality and Host. Berkeley (University of California Botanical Gardens), Alameda County, California, on horse crippler cactus, Homalocephala texensis (Cactaceae). North American Distribution. California, Mexico, New Mexico, and Texas. Additional Hosts. The principal hosts of this mealybug belong to the Cactaceae. A few of the host species of this family are recorded as follows: Cereus formosus monstrosus, C. peruvianus, Echinomastus johnsonii, Echinopsis sp., Hamatocactus setispinus, Mammillaria macdougalii, Mammillaria spp., Myrtillocactus geometriazans, and Wilcoxia sp. External Features and Habitat. At full maturity the female of this species is evenly covered with a short, light-gray, waxy secretion. The lateral filaments are lacking, and the caudal ones very short and inconspicuous. A definite ovisac is produced at maturity (see col. pi. II). When specimens are boiled in lacto-phenol or potassium hydroxide during the mounting procedure, they turn dark and much of the body contents often form into large blue-black, round globules which are usually quite difficult to remove without damage to the integument. The mealybug occurs exposed on the green, fleshy vegetation and spines, as well as around the crown of its cactus host. Recognition Characters. Adult females, mounted, 2.10 to 3.20 mm long, 1.35 to 2.25 mm wide; body form broadly oval. Dorsum with cerarii rather poorly developed, there being normally 7 pairs on the abdomen, and from 2 to 7 pairs on thorax and head, these difficult to discern because cerarian setae are usually much more slender and generally situated farther apart, total recognizable cerarii running as high as possibly 14 or 15 pairs. Anal lobe cerarius with 2 rather small, slender conical setae, 3 or 4 slender auxiliary setae of various lengths, very few scattered trilocular pores, no sclerotization; remaining cerarii along abdominal margin with but 2 conical setae, these normally more slender and, at times, more widely separated than anal pair, no auxiliary setae except possibly in certain cerarii on head and thorax, scarcely any concentration of trilocular pores. Trilocular pores evenly distributed over entire dorsum. Minute circular pores numerous, scattered over dorsal surface. Tubular ducts of oral-collar type absent. Oral-rim tubular ducts comparatively few, arranged in a single row of from 6 to 8 across each abdominal segment, except the last, few scattered on thoracic segments and head. Body setae all small and slender. Anal ring of normal form and size for genus; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores considerably numerous, situated predominantly in midregion of abdomen, from posterior apical to fourth segment, absent elsewhere on venter. Trilocular pores generally

distributed over venter. Minute circular pores numerous, scattered over ventral surface. Small oral-collar tubular ducts occur in same general area as multiloculars, and in considerable numbers in lateral areas of eighth forward to third abdominal segments, very few in lateral area of metathorax, absent elsewhere on venter. Oral-rim tubular ducts of same size as those on dorsum, occur in small numbers submarginally on abdomen, thorax and head, very few scattered in sternal area. Body setae slender, generally longer than those on dorsum. Circulus quite large at times, faintly defined in fully developed females, usually appearing broadly-oval, capable of folding along intersegmental line between which it lies. Legs well developed, fairly small for members of this genus; hind coxae with cluster of translucent pores at base; apical half of hind tibiae with small irregular cluster or patch of translucent pores; claw with very small denticle on plantar surface. Antennae 8-segmented, slender, eighth segment often slightly constricted near tip to form small projection. Notes. This species is perhaps most closely related to Spilococcus quercinus McKenzie but differs from it mainly in possessing far fewer dorsal oral-rim tubular ducts on abdominal segments, each entire segment except last, with less than 10 such ducts; 14 or 15 pairs of cerarii; and normally a cactus host. S. quercinus, on the other hand, shows 20 or more oral-rim tubular ducts on each abdominal segment, except last; usually not more than 11 discernible cerarian pairs; and occurs on oak and Ceanothus. California Records (Map 160) Alameda Co.: Berkeley (U. C. Botanical Gardens), III—25—58, on "cactus" (W. S. Sibray, CDA); Berkeley, IV-11-60, on "cactus" (W. S. Sibray and T. R. Haig, CDA); 1-24-^9, on Cereus formosus montrosus (W. S. Sibray, CDA); Berkeley (U. C. Botanical Gardens), XII-1948, on Echeveria longissima (R. E. Beer, USNM); 1-20-59, on Echinomastus johnsonii (H. L. McKenzie, UCD); XI-4-58, on Hamatocactus setispinus and Wilcoxia sp. (A. E. Pritchard and H. L. McKenzie, UCD); X-4-48 and X - l l - 4 8 , on Homalocephala texensis (R. E. Beer, USNM); X-ll—48, on Mammillaria dolichocentron (R. E. Beer, USNM); 1-24-49, on Mammillaria macdougalii (W. S. Sibray, CDA); Hayward, VII10-50, on "cactus" (W. S. Sibray, CDA); 111-19-62, on "cactus" (P. Sweigart, CDA); 1-28-49, on Echinopsus sp. (W. S. Sibray, CDA); 1-31-49, on Echinopsus multiplex (W. S. Sibray, CDA); 1-31-49, on Myrtillocactus sp. (W. S. Sibray, CDA); Oakland, II-1-49, on Cactus sp. (W. S. Sibray, CDA); 1-29-49, on Echinopsus sp. (W. S. Sibray, CDA); II—1—49, on Lophophora williamsi (G. B. Laing, CDA). Contra Costa Co.: Richmond, X-23-41, on Myrtillocactus geometrizans (M. R. Bell, CDA?). Los Angeles Co.: Baldwin Park, 1-9-^40, on "cactus" (H. L. Marsh, LEM); Glendale, VII-20-42, on "cactus" (L. E. Myers, LEM); Hynes (nursery), 1-1945, on Piptanthocereus sp. (Boerner, LEM); La

TAXONOMY Canada, III-7-57, on "cactus" (Hodge, CDA); Lennox, VII-9-41, on Mammillaria sp. (F. R. Piatt, LEM); Los Angeles (nursery), VI-26-41, on "cactus" (V. E. Daniels, LEM); Los Angeles VI-26-41, on "cactus" (H. A. Robinson, UCD); Los Angeles (nursery), VII-22-40, on Cereus peruvianus (F. R. Piatt, L E M ) ; V I I I - 1 8 ^ 1 , on Echinopsus silvestrii (F. R. Piatt, LEM); IV-9-45, on Homalocephala texensis (R. H. Smith, UCD); Paramount, XI-13-50, on "cactus" (French, UCD) ; 1-18-62, on Coryphantha andrea (J. Wood and J. Weber, CDA); Pasadena, VIII-14-41, on "cactus" (V. E. Daniels, LEM); Pomona, IX-29-38, on "cactus" (Vandenberg, LEM); Pomona (nursery), 1-20-43, on "cactus" (V. E. Daniels, LEM) ; West Los Angeles (nursery), VII-29-41, on Myrtillo sp. (J. Caldwell, LEM). Orange Co.: Anaheim, XII-13-63, on Haworthia margaritijera (R. Wyatt, CDA). Sacramento Co. : Sacramento, XI-23-62, on "cactus" (M. Scribner, CDA). San Diego Co.: Fallbrook, XI-27-63, on Astrophytum sp. (M. K. Hess and H. T. Oldham, CDA); San Diego, IX-15-60, on "cactus" (J. R. Carlin, CDA); VII-27-61, on "cactus" (Beai and Knott, CDA). San Francisco Co.: San Francisco, XI-20-56, on "cactus" (R. Bozzini and D. J. Bingham, CDA); 1-28-42, on "cactus" (collector ?, CDA); 11-3^9, on Mammillaria sp. (A. L. Seeley and S. M. Mather, CDA). San Mateo Co. : South San Francisco, II-6-62, on "cactus" (B. Davis, CDA). Sonoma Co.: Petaluma, XII-4-41, on Mammillaria sp. (M. R. Bell, CDA).

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Spilococcus ceanothi McKenzie, new species Squawcarpet Mealybug (Figure 171) Type Locality and Host. Type and paratype adult females on Squawcarpet or Mahala Mat, Ceanothus prostratus (Rhamnaceae) (crown and roots), 13 miles east Nevada City, Nevada County, California, collected October 1, 1964, by D. R. and J. F. Miller. Additional paratype adult females on the same host, Ceanothus prostratus, at Nevada City, Nevada County, California, collected July 7, 1964, by E. Remmers; on same host, 3 miles east Canyondam, Plumas County, California, collected July 9, 1964, by D. R. Miller; on the same host, 2 miles north Mt. Shasta, Siskiyou County, California, collected November 9, 1965, by T. R. Haig; and on Ceanothus pumilus (roots), 6 miles northwest Weaverville, Stoney Creek on Trinity Lake, Trinity County, California, collected October 11, 1963, by G. M. Buxton. Type Material. Holotype adult female (single specimen on slide) and female paratypes in museum, University of California, Davis. Female paratypes in United States National Collection of Coccoidea (U. S. National Museum), Washington, D. C., and in collection of California State Department of Agriculture, Sacramento. North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. This mealybug is covered with a heavy, white, fluffy, wax secretion. The body form is subcircular; body color gray to light brown; legs red. There is 1 pair of thin, minute, caudal filaments. A filamentous ovisac is produced which covers all but the head of the insect. It is 3 to 5 times longer than length of normal adult. The mealybug is found on the decumbent branches, crown, and occasionally roots of its host. Ants were observed tending this mealybug. Recognition Characters. Adult females, mounted, 2.80 to 3.00 mm long, 2.00 to 2.40 mm wide; body form broadly oval to rotund. Dorsum with cerarii somewhat variable in number, not more than 15 pairs, including 8-9 recognizable pairs on abdomen and metathorax, 1-4 thoracic pairs, 1-3 head pairs, these often represented by a single conical seta. Anal lobe cerarius with 2 rather slender conical setae, 3-4 more slender auxiliary setae, 1 of which is often quite thick and long, few obviously associated trilocular pores. Remaining abdominal cerarii each with 2 conical setae, progressively slightly smaller with distance from anal lobes, no auxiliary setae, slight concentration trilocular pores; cerarii along thoracic margin similarly constructed when present, although at times conical setae widely separated, or represented by only a single seta; frontal and ocular cerarii usually present, each of these with 2 or 3 small conical setae, no slender auxiliary setae, slight concentration trilocular pores; postocular cerarii, when present, usually represented by a single conical seta.

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Fig. 171. Spilococcus ceanothi McKenzie, new species, collected on crown and roots of squaw carpet, Ceanothus prostratus (Rhamnaceae), 13 miles east Nevada City, Nevada County, California.

Sitlriyeu Dorsum evenly beset with trilocular pores; less numerous minute circular pores. Oral-rim tubular ducts very sparse, a broken series of 2 or 3 on each side along submargin of abdomen, absent on thorax and head. A few oral-collar tubular ducts along margin of last 4 or 5 abdominal segments. Body setae short and slender. Anal ring slightly removed from posterior margin of ninth abdominal segment, normal for genus; each of its 6 setae about twice as long as diameter of ring. Venter with multilocular disk pores present from posterior apical segment to fifth abdominal segment in midregion of abdomen, absent elsewhere on venter. Trilocular pores fairly evenly beset on venter, except for certain "clear" areas on thorax and head. Minute circular pores scattered on venter. Oral-rim tubular ducts occur in limited numbers along submargin of third abdominal segment forward to thorax and head, absent elsewhere on venter. Oral-collar tubular ducts fairly numerous on abdomen, scattered on thorax, absent on head. Most body setae longer than those on dorsum. Circulus rather large, irregularly quadrate, apparently located on fourth segment, not divided by segmental line. Legs comparatively short and stout; hind coxa with numerous translucent pores at base; tibia with irregular clusters of similar pores, especially noticeable at distal half of segment; slight denticular swelling on plantar surface of claw. Antennae 7-segmented. Notes. This species seems rather closely related to Spilococcus andersoni (Coleman), but differs in having a 7-segmented antenna; fewer ventral submarginal oral-rim tubular ducts on third abdominal segment forward to prothorax; and a much different host, Ceanothus spp., especially C. prostratus (Rhamnaceae). Spilococcus andersoni, on the other hand, possesses an 8-segmented antenna; fairly numerous ventral submarginal oral-rim tubular ducts on third abdominal segment forward to prothorax; and its hosts species are mainly representatives of the Cupressaceae, Taxaceae, and Taxodiaceae. California Records (Map 161) Nevada Co.: 13 miles east Nevada City, X - l - 6 4 , on Ceanothus prostratus (D. R. and J. F. Miller, UCD) ; Nevada City, Scott's Flat Road, VII-7-64, on Ceanothus prostratus (E. Remmers, CDA). Plumas Co.: 3 miles east Canyondam, VII-9-64, on Ceanothus prostratus (D. R. Miller, U C D ) . Siskiyou Co. : 2 miles north Mt. Shasta, X I - 9 - 6 5 , on Ceanothus prostratus (T. R. Haig, C D A ) . Trinity Co. : 6 miles northwest Weaverville, Stoney Creek on Trinity Lake, X - l l - 6 3 , on Ceanothus pumilus (roots) (G. M. Buxton, C D A ) .

Spilococcus corticosus McKenzie, new species Bark Mealybug (Figure 172) Type Locality and Host. Type and paratype adult females on common sagebrush, Artemisia tridentata (Compositae), Warner Springs, San Diego County,

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California, collected January 26, 1965, by D. R. Miller. Additional paratype adult females on California sagebrush, Artemisia californica (Compositae), Whittier, Los Angeles County, California, collected October 21, 1912, by P. H. Timberlake. Type Material. Holotype adult female (single specimen on slide) and female paratypes in museum, University of California, Davis. North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. The adult female of this species is covered with a smooth, white, waxy secretion. The body color is green; body shape oval; legs red. A filamentous ovisac is produced which encloses the entire female body; the eggs are yellowishgreen. This mealybug is found above ground beneath the bark of its host. No ants were found in attendance. This mealybug occurs under the bark near the crown of its host. Recognition Characters. Adult females, mounted, 2.80 to 3.00 mm long, 1.20 to 1.50 mm wide; body form broadly oval. Dorsum with cerarii extremely variable, often represented by a single conical seta, with as few as 5 and not more than 14 pairs, including 3 - 7 pairs on abdomen, 0 - 5 thoracic pairs, and 2 - 3 head pairs. Anal lobe cerarius with 2 conical setae, 3 - 5 slender auxiliary setae, few associated trilocular pores. Remaining abdominal cerarii each with 2 conical setae, progressively slightly smaller with distance from anal lobes, no auxiliary setae, slight concentration trilocular pores; cerarii along thoracic margin similarly constructed when present, although at times conical setae widely separated, represented by only a single seta, or at times 3 conical setae; frontal and ocular cerarii

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Fig. 172. Spilococcus corticosus McKenzie, new species, collected on common sagebrush, Artemisia tridentata (Compositae), at Warner Springs, San Diego County, California.

usually present, each with 2 or 3 small conical setae, no auxiliary setae, slight concentration trilocular pores. Dorsum evenly beset with trilocular pores; less numerous minute circular pores. Oral-rim tubular ducts in a broken submarginal series of single pores from eighth abdominal segment forward to head, very few scattered elsewhere on abdomen and thorax. Oral-collar tubular ducts of 2 distinct sizes numerous, situated in irregular transverse bands across abdominal segments, in submarginal clusters on thorax, scattered in median area of thorax and head. Body setae short and slender. Anal ring apical, normal for genus; each of its 6 setae about twice as long as diameter of ring itself. Venter with multilocular disk pores present from last segment of abdomen forward to posterior margin of fifth segment, infrequently 1 or 2 such pores on head. Trilocular pores fairly evenly beset on venter, except for certain "clear" areas on thorax and head. Minute circular pores scattered on venter. Oral-rim tubular ducts usually absent on venter, infrequently a single submarginal one evident on third or fourth abdominal segments. Oral-collar tubular ducts of 2 sizes, similar to those on dorsum, segmentally arranged on abdomen, tending to cluster along submargin of these segments, in similar clusters along submargin of thorax, noticeable group opposite midand hind coxae, anterior spiracle and body margin, scattered elsewhere on thorax and head. Most body setae longer than those on dorsum. Circulus present or absent (2 specimens with a circulus, 1 without this structure); when present, quite small, oval, apparently divided by segmental line across which it lies. Legs comparatively short though slender; hind coxa with numerous translucent pores at base; apparently these structures absent on tibiae; slight denticular swelling on plantar surface near tip of claw. Antennae 8-segmented. Notes. Specimens of this species which possess a circulus seem rather closely related to Spilococcus cactearum McKenzie, but differ principally in having numerous dorsal oral-collar tubular ducts, especially noticeable on abdomen; in the species last mentioned, these structures are apparently absent on dorsal surface of body. One specimen of S. corticosus lacks a circulus. It resembles S. keiferi McKenzie in possessing a cluster of ventral oral-collar tubular ducts situated between anterior spiracle and body margin, but differs from the species last indicated in possessing about 14 pairs of cerarii, some of these occurring on thorax and head; and numerous dorsal oral-collar tubular ducts. Spilococcus keiferi, on the other hand, has but 6 or 7 recognizable pairs of cerarii on abdomen counting forward from anal lobes, and none on thorax and head; only a very few oral-collar tubular ducts scattered over dorsal body surface. California Records (Map 162) Los Angeles Co.: Whittier, X - 2 1 - 1 2 , on Artemisia californica (P. H. Timberlake, U C D ) . San Diego Co.: Warner Springs, 1-26-65, on Artemisia tridentata (D. R. Miller, U C D ) .

Spilococcus er logorìi (Ehrhorn) Eriogonum Mealybug (Figures 173 and 174) Dactylopius eriogoni Ehrhorn, 1899. Canad. Ent. 31:103. Erium eriogoni (Ehrhorn), Cockerell, 1902. Ann. Mag. Nat. Hist. 10(7):465. Erium eriogoni (Ehrhorn), Fernald, 1903. Mass. Agr. Expt. Sta. Spec. Bull. 88, p. 112. Pseudococcus yerba-santae Essig, 1913. Jour. Ent. and Zool. 5 ( 2 ) : 8 5 - 8 7 , illus. Pseudococcus eriogoni (Ehrhorn), Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., p. 44. Spilococcus eriogoni (Ehrhorn), Ferris, 1950. Atlas of scale insects of North America, Ser. V : 2 2 3 224, illus. Spilococcus atriplicis (Cockerell), Ferris, 1953. Ibid., Vol. VI: 446 (eriogoni synonymized with atriplicis) . Spilococcus eriogoni (Ehrhorn), McKenzie, 1960. Hilgardia 2 9 ( 1 5 ) :761 (eriogoni recognized as distinct from atriplicis). Spilococcus haigi McKenzie, 1962. Hilgardia 3 2 ( 1 4 ) : 6 7 9 - 6 8 3 , illus. (New synonymy). Type Locality and Host. Stevens Creek Canyon, Santa Clara County, California, on Eriogonum sp. (Polygonaceae). North American Distribution. California, Maryland, Mexico, and Texas. Additional Hosts. Some of the hosts attacked by this mealybug include: Artemisia californica (Compositae), Asparagus sp. (Liliaceae), Compositae, Cressa eretica (Convolvulaceae), Eriodictyon californicum, E. tomentosum (Hydrophyllaceae), Eriodictyon spp. (Hydrophyllaceae), Eriogonum fas-

424

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Fig. 173. Spilococcus eriogoni (Ehrhorn), collected on Eriogonum sp. (Polygonaceae), at Stevens Creek Canyon, Santa Clara County, California.

TAXONOMY

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Fig. 174. Spilococcus eriogoni (Ehrhorn), collected on Malacoihrix sonchoides (Compositae), at Fernley, Lyon County, Nevada. Note especially the presence of a weekly developed circulus, a structure normally absent in this species. Certain California specimens demonstrate this type of circulus, although all other structures are quite typical for the species.

425

426

TAXONOMY

OF

SPECIES

ciculatum, E. inflatum, E. latifolium var. saxicola (Polygonaceae), Eriogonum spp. (Polygonaceae), Gossypium sp. (Malvaceae), Gutierrezia californica (Compositae), Heterotheca sp. (Compositae), Lathy rus littoralis (Leguminosae), Parosela (= Dalea) emoryi (Leguminosae), Solidago sp. (Compositae), and Suaeda sp. (Chenopodiaceae). External Features and Habitat. According to the original description, the adult female of the species is "enclosed in a densely woven white felt sac about 2Vi mm long and 1 mm broad; also secreting considerable loose cottony matter." This mealybug occurs on the roots, crown, stems, and foliage of the host. Recognition Characters. Adult females, mounted, 1.50 to 3.00 mm long, 1.00 to 2.10 mm wide; body form broadly oval. Dorsum with number of pairs of cerarii reduced, there being 7-8 that are recognizable, counting forward from anal lobes, at times an ocular or frontal pair present, along thoracic margins the setae become so small and widely separated that distinct cerarii are unrecognizable. Anal lobe cerarius with 2 rather small and slender conical setae. 3-4 slender auxiliary setae, few scattered trilocular pores. Remaining cerarii with but 2 setae, no auxiliary setae, scarcely any concentration trilocular pores. Trilocular pores fairly numerous, scattered over entire dorsum. Minute circular pores scattered. Tubular ducts of oral-collar type absent. Oral-rim tubular ducts numerous, arranged in a single row of as many as 20 across each abdominal segment, except the last, scattered over thoracic segments and head. Body setae small and slender. Anal ring apical, of normal form and size; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores situated in midregion on apical segment forward to posterior margin of fifth abdominal segment, absent elsewhere on body. Trilocular pores fairly numerous, perhaps somewhat less abundant in sternal area. Minute circular pores scattered. Oral-collar tubular ducts occur in same area as multilocular disk pores, also in considerable numbers in lateral areas of eighth to sixth and possibly fifth segments. Oral-rim tubular ducts of same size as those on dorsum occur in small numbers in lateral areas of all segments from eighth abdominal forward to head, in midregion of abdominal segments anterior to sixth, in sternal areas, and fairly numerous about spiracles. Body setae slender, some noticeably longer than those of dorsum. Circulus absent, or rarely present. Legs well developed; hind coxa with translucent pores in small group near distal end of segment; claw usually with small denticle on plantar surface. Antennae normally 8-segmented, rarely 7-segmented, the eighth segment at times showing tendency to be divided. Notes. Ferris (1953) synonymized this species with Spilococcus atriplicis (Cockerell). On the bases of an examination of a considerable amount of material from various hosts and localities in California,

believed to represent S. eriogoni, it is the author's (1960) opinion that the 2 species are definitely distinct. This statement is based on the fact that S. eriogoni normally has a reduced number of cerarii, there being only 7 to 8 pairs counting forward from the anal lobes, and at times a recognizable ocular or frontal pair present, whereas in S. atriplicis there are at least 15 recognizable pairs of cerarii. In addition, S. eriogoni usually possesses almost twice as many dorsal oral-rim tubular ducts, particularly on the fifth and sixth abdominal segments, as compared to S. atriplicis. In view of this development, the author believes Ferris' opinion as expressed in his 1953 key to species, where he actually separated S. atriplicis and S. eriogoni, is sounder than his statement under the discussion of S. atriplicis that the 2 are identical. Spilococcus eriogoni also has a tiny denticle on plantar surface of claw, this feature not included on drawing of species in Ferris' Atlas. In this study, Spilococcus haigi McKenzie, a species originally described from Nevada, is placed as a synonym of S. eriogoni. This came about as a result of examination of numerous collections of S. eriogoni, certain specimens of which possessed a circulus. In all other respects these examples are identical to genuine S. eriogoni. California Records (Map 163) Alameda Co.: Fremont, IX-2-62, on Eriogonum sp. (T. R. Haig and D. A. Taylor, CDA). Fresno Co.: Panoche Pass, 111-30-39, on Aplopappus sp. (G. F. Ferris, UCD). Humboldt Co.: 1 mile south Samoa, VIII-11-63, on Lathyrus littoralis (foliage) (D. R. Miller and J. S. Buckett, UCD). Imperial Co.: Bard, IV-6-53, on Gossypium sp. (E. J. Dietrick, CDA); Calexico, XII-23-39, on Asparagus sp. (Mexican) (C. G. Anderson and A. J. Hansen, CDA); 8 miles west Glamis, XII-23-65, on Eriogonum deserticola (D. R. and J. F. Miller, UCD). Kern Co.: 11 miles west Randsburg, X-22-55, on "weed" (J. W. Stockton, CDA). Los Angeles Co.: 20 miles south Gorman, VII28-63, on Compositae (crown and roots—associated with ants, Crematogaster coarctata vermiculata) (D. R. Miller, UCD). Modoc Co.: 5 miles south Cedarville, VI-28-63, on Eriogonum latifolium var. saxicola (roots) (D. R. Miller, UCD). Orange Co.: Atwood, X-5-64, on Heterotheca sp., tended by Solenopsis xyloni (D. H. Byers, CDA); 1 mile south of South Laguna, VII-26-63, on Artemisia californica (roots) (D. R. Miller, UCD). Riverside Co.: 3 miles west Garnet, II-5-60, on Compositae (U. Nur, UCD); 3.4 miles northeast Gilman Springs, III—9—63, on Gutierrezia californica (D. R. Miller, UCD); Riverside, X-2-33, on ? (S. E. Flanders, USNM); 1-30-36, on Croton californicus (J. D. Maple, UCD); Thermal, XII-6-56, on Eriogonum inflatum (R. P. Allen, CDA). San Diego Co.: Del Mar, VII-26-12, on Eriodictyon tomentosum (P. H. Timberlake, UCD); Encini-

TAXONOMY tas, VI-13-60, in "soil" (H. L. McKenzie, UCD); Nallecitos, XII-23-39, on Suaeda sp. (R. R. McLean, CDA). San Joaquin Co.: Escalon, IX-5-31, on "spike weed" (T. Heurlin, USNM). Santa Clara Co.: Los Altos (Hidden Villa Road), VIII-23-16, on Eriogonum sp. (G. F. Ferris, USNM); Los Altos (Adoba Gulch), VII-25-16, on Eriogonum sp. nudum ? (G. F. Ferris, UCD); Stevens Creek Canyon (near Mt. View), 1898 ?, on Eriogonum sp. (E. M. Ehrhorn, USNM); Stevens Creek Road, V-5-17 and V-6-17, on Eriogonum sp. (roots) (G. F. Ferris, UCD). Siskiyou Co.: 1 mile south Hornbrook, VIII11-64, on Eriogonum sp. (D. W. Robinson and T. R. Haig, CDA); 2 miles north Mt. Shasta, VIII16-65, on Eriogonum latifolium var. nudum (T. R. Haig, CDA). Stanislaus Co.: Near Modesto (Puerto Canyon), III—1935, on Eriogonum fasciculatum (G. F. Ferris, UCD). Trinity Co.: 5 miles north Burnt Ranch, VIII12-63, on Eriogonum latifolium (crown and roots— associated with ants, Tapinoma sessile) (D. R. Miller, UCD). Ventura Co.: Fillmore, VI-24-36, on Croton californicus (collector ?, UCD); Oxnard, IX-30-55, on Cressa cretica (C. May, CDA); Sespe (Sespe Canyon), V - l l - 1 1 , on Eriodictyon californicum (E. O. Essig, CAS); Moorpark, 11-27-36, on Croton californicus (J. D. Maple, UCD). Yolo Co.: Rumsey, XII-19-64, on Eriogonum sp. (D. R. and J. F. Miller, UCD).

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Spilococcus geraniae (Rau) Geranium Mealybug (Figure 175) Trionymus geraniae Rau, 1938. Canad. Ent. 70:162165, illus. Spilococcus geraniae (Rau), McKenzie and Williams, 1965. Hilgardia 37(1):7-9, illus. Type Locality and Host. Saratoga Springs, Saratoga County, New York, on geranium, Geranium robertianum (Geraniaceae). North American Distribution. California and New York. Additional Hosts. Artemisia douglasiana (Compositae). External Features and Habitat. According to the original description, this species has the wax evenly distributed over the body, except between body segments where the color appears to be greenish. Specimens with waxy secretion removed are, therefore, greenish, and specimens freshly killed in alcohol turn green. A long and well-developed ovisac is produced which arises from the underside of the posterior part of the body. This mealybug lives habitually alone, exposed on the underside of the leaves or in the axils where the leaves join the main stem of its host. Recognition Characters. Adult female, mounted, approximately 2.50 mm long, 1.40 mm wide. Dorsum with 6 or 7 pairs of abdominal, and possibly 1 or 2 pairs of frontal, cerarii. Anal lobe cerarius without sclerotization, with a pair of stout conical setae, 3 or 4 slender auxiliary setae, and surrounded by a cluster of trilocular pores. Remaining abdominal cerarii each with 2 conical setae (progressively smaller with distance from anal lobes), without auxiliary setae, but with a few trilocular pores. Cerarii on head, if correctly interpreted, with 2 or 3 slender conical setae and a few trilocular pores. Trilocular pores evenly distributed on dorsum. Oral-collar tubular ducts present along lateral margin from anal lobes forward to fourth abdominal segment; absent elsewhere on dorsum. Oral-rim tubular ducts in transverse bands of from 6 to 14 on each abdominal segment except the last, also scattered over thorax and head. Body setae short and slender. Anal ring apical, with no unusual characters; each of its 6 setae about twice as long as diameter of ring. Venter with multilocular disk pores abundant, in transverse, double-row bands from apex to posterior margin of fifth abdominal segment, a few submarginal ones present along posterior margin of fourth segment; and often with 1 or 2 pores in prothoracic region between anterior coxa and body margin. Trilocular pores evenly distributed. Oral-collar tubular ducts small, all essentially the same size, fairly numerous, in tranverse bands on abdominal segments, and a conspicuous cluster on prothorax between anterior coxa and body margin. A few oral-rim tubular ducts of the same size as those on dorsum occur between anterior and posterior spiracles and body margin, 1

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Fig. 175. Spilococcus geraniae (Rau), collected on geranium, Geranium robertianum (Geraniaceae), Saratoga Springs, Saratoga County, New York (paratype). The single adult female collected on Artemisia douglasiana (Compositae) at Sacramento, California agrees in detail with this illustration.

TAXONOMY

or 2 on head. Body setae slender, slightly longer than those on dorsum. Circulus small, transversely oval, not divided by segmental line. Legs well developed, comparatively large; a few rather indistinct translucent pores on hind coxa; a few similar pores distributed on femora and tibiae of middle and hind legs; claw without denticle. Mouthparts comparatively broad. Antennae 8-segmented. Eyes present. Notes. This species is closely related to Spilococcus parvicirculus McKenzie. Spilococcus geraniae differs in these respects: very numerous ventral multilocular disk pores in double rows on the abdomen (approximately 500); a conspicuous cluster of ventral oralcollar tubular ducts occurs between the anterior coxa and the body margin; and the ventral oral-rim tubular ducts are absent on the abdomen. On the other hand, S. parvicirculus has fewer ventral multiloculars in single rows on the abdomen (approximately 140); the oral-collar tubular ducts are absent on the thorax; and the ventral oral-rim tubular ducts are present along the submargin of the abdomen. A single immature specimen of Spilococcus geraniae (Rau) was found by W. R. Bauer, California State Department of Agriculture, April 8, 1966 on foliage of Artemisia douglasiana (Compositae) growing along the banks of the American River in Sacramento, California. Artemisia plants were taken into the laboratory for propagation. The mealybug was reared to maturity on these plants, was subsequently killed and ultimately mounted on a microscope slide June 20, 1966. The single example agrees in detail with paratype specimens of S. geraniae, from which the accompanying illustration was made. California Records (Map 164) Sacramento Co.: Sacramento (along American River), VI-20-66, on Artemisia douglasiana (W. R. Bauer, CDA).

Map 164. Distribution in California of Spilococcus geraniae (Rau).

SattBmd$fSmMmtto

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Spilococcus gutierreziae (Cockerell) Gutierrezia Mealybug (Figure 176) Dactylopius gutierreziae Cockerell, 1896. Jour. N. Y. Ent. Soc. 4:203. Pseudococcus gutierreziae (Cockerell), Fernald, 1903. Mass. Agr. Expt. Sta. Spec. Bull. 88, p. 103. Spilococcus gutierraziae (Cockerell), Ferris, 1950. Atlas of scale insects of North America, Ser. V:225-226, illus. Type Locality and Host. Mesilla Valley between Las Cruces and Organ, Dona County, New Mexico, on matchweed, Gutierrezia sarothrae (Compositae). North American Distribution. Arizona, California, New Mexico, Texas, and Utah. Additional Hosts. Haplopappus sp. (Compositae). External Features and Habitat. According to the original description, the adult female of this mealybug is slate-colored, and forms a long, firm, snow-white ovisac, 4.00 to 6.00 mm long and 1.00 mm wide. This species occurs on the leaves of its host. Recognition Characters. Adult females, mounted, 1.90 to 2.00 mm long, 1.00 to 2.00 mm wide; body form broadly oval. Dorsum with 15 or 16 pairs of cerarii; anal lobe cerarius with 2 small, slender conical setae, 1 or more elongate slender auxiliary setae, very small concentration trilocular pores; remaining cerarii with 2 conical setae, no auxiliary setae, and practically no concentration trilocular pores-—setae becoming so small along lateral margins of thorax that cerarii can scarcely be recognized; frontal cerarius seems to be entirely absent. Trilocular pores quite evenly distributed on dorsum. Oral-collar tubular ducts absent. Oral-rim tubular ducts of 2 distinct sizes on dorsal surface; smaller ones numerous, occurring on eighth abdominal segment forward to posterior border of mesothorax; larger ones less numerous, situated from mesothorax to head. Body setae short and slender. Anal ring small and compact; each of its 6 setae only slightly longer than greatest diameter of ring. Venter with multilocular disk pores present in small numbers from posterior apical segment forward to fifth abdominal segment, confined to midregion, also in lateral areas of seventh and eighth segments. Trilocular pores not as numerous as on dorsum, rather large "clear" areas appearing especially in sternal area where these pores are absent. Oral-collar tubular ducts extremely small, few, situated in same general area as multilocular disk pores. Oral-rim tubular ducts of a slightly smaller size than those on dorsum present in anterior portion of abdominal midregion and thoracic areas, and in lateral areas of abdomen forward to mesothorax; larger sized oral-rim tubular ducts present in small numbers in extreme lateral areas of mesothorax and prothorax. Body setae sparsely distributed, some noticeably longer than those on dorsum.

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Fig. 176. Spilococcus gutierreziae (Cockerell), collected on Gutierrezia sp. (Compositae), at Deep Springs Valley, Inyo County, California. (Illustration prepared from specimens collected on Gutierrezia sp. [Compositae], El Paso, El Paso County, Texas.)

TAXONOMY Circulus absent. Legs well developed, though noticeably small; hind coxa with few small translucent pores; tibia with very few translucent pores situated on distal half of segment; claw without denticle on plantar surface. Antennae 8-segmented. Notes. This mealybug is readily distinguishable from other species of this group lacking a circulus by the presence of 2 distinct sizes of oral-rim tubular ducts, those on dorsum anterior to metathorax and on head distinctly larger than the ones situated in abdominal area forward to mesothorax. The species is, apparently, not too commonly found in California. California Records (Map 165) Imperial Co.: Mountains Springs, VI-21-64, on Haplopappus cooperi ? (D. R. Miller, U C D ) . Inyo Co.: Deep Springs Valley, VII-1918, on Gutierrezia sp. (G. F. Ferris, U C D ) . Kern Co.: Monolith, IX-17-66, on Gutierrezia sarothrae (T. R. Haig, C D A ) .

Spilococcus implicatus Ferris Implicated Mealybug (Figures 177 and 178; Color Plates III and X X I V ) Spilococcus implicatus Ferris, 1950. Atlas of scale insects of North America, Ser. V:227-228, illus. Type? Locality and Host. Stanford University, Santa Clara County, California, on redwood, Sequoia sempervirens (Taxodiaceae). North American Distribution. California and Illinois. Additional Hosts. Cupressus macrocarpa and Cupressus spp. (Cupressaceae), Juniperus sp. (Cupressaceae), and Matthiola sp. (Cruciferae).

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External Features and Habitat. The adult female of this mealybug is covered with a dull gray, waxy secretion, without lateral filaments, caudal ones slightly developed, very short, or obsolete. A long, firm ovisac is produced (see col. pis. Ill and X X I V ) . This species occurs entirely on the foliage of its host. Recognition Characters. Adult females, mounted, 1.95 to 2.50 mm long, 1.20 to 1.70 mm wide; body form broadly oval. Dorsum with cerarian pairs quite variable in number, total ranging from 7-15, with 4 - 8 pairs on abdomen, 3 - 7 pairs on anterior thorax and head; anal lobe cerarius with 2 rather slender conical setae, 3 - 4 slender auxiliary setae, very few trilocular pores; anterior to anal lobes on abdomen, cerarii each with 2 small conical setae, without auxiliary setae, practically no concentration trilocular pores. Trilocular pores evenly beset on dorsum. Oralcollar tubular ducts along submargin of posterior abdominal segments. Oral-rim tubular ducts very few, 1 to 3 situated on each side along margin of head, 0 to 4 or 5 on each side along margin of posterior abdominal segments, absent elsewhere on dorsum. Body setae small and slender. Anal ring apical, normal in size and form for genus; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores in midregion of abdomen from posterior apex forward to fourth segment. Trilocular pores not as numerous as on dorsum, rather "clear" areas evident especially in sternal area where these pores are absent. Minute circular pores scattered. Oral-collar tubular ducts rather numerous, often in clusters, in lateral areas of abdominal and thoracic segments. Oral-rim tubular ducts similar in size to those on dorsum, occur in a conspicuous submarginal band on abdomen, and in postspiracular area of thorax (Ferris' [1950] illustration of this species shows these structures also present in sternal areas of metathorax and mesothorax). Body setae sparsely distributed, some noticeably longer than those on dorsum. Circulus absent. Legs large, and well developed; hind coxa with small translucent pores; hind tibia without translucent pores; claw without denticle on plantar surface. Antennae 8-segmented. Notes. According to Ferris (1950), this species was included on the type slides of Spilococcus sequoiae (Coleman), and has been entangled with the latter, and probably also with Spilococcus andersoni (Coleman), ever since. It is probably most closely related to S. sequoiae, but differs from it mainly in the absence of ventral lateral clusters of multilocular disk pores on abdomen, these structures quite obvious in S. sequoiae. It is easily separated from other known species of this genus lacking a circulus by the absence of numerous dorsal oral-rim tubular ducts, only a very few of these structures evident along body margins of head and posterior abdominal segments, at times absent. Spilococcus implicatus seems quite variable in certain morphological structures. Specimens collected on

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Fig. 177. Spilococcus implicatus Ferris, collected on redwood, Sequoia (Taxodiaceae), at Stanford University, Santa Clara County, California.

sempervirens

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\\\i D

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Fig. 178. Spilococcus implicatus Ferris, collected on Monterey cypress, Cupressus macrocarpa (Cupressaceae), Shell Beach, San Luis Obispo County, California. Note especially the more numerous cerarii, and extra ventral oral-rim tubular ducts between anterior spiracle and body margin.

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Cupressus sp. and Matthiola sp. at Camarillo, Ventura County, California show more dorsal oral-rim tubulars, and those taken on Cupressus macrocarpa at Shell Beach, San Luis Obispo County, California possess more cerarii than is depicted in Ferris' (1950) illustration of the species. California Records (Map 166) California, X-1897, on Cupressus macrocarpa (A. Craw, USNM). Alameda Co.: Berkeley, VIII-15-35, on Cupressus sp. (G. B. Laing, C D A ) ; 1-23-63, on Juniperus sp. (R. Cruickshank, C D A ) ; XII-14-34, on Cupressus macrocarpa (D. W. Clancy, U C D ) . Contra Costa Co.: El Cerrito, VI-23-59, on Cupressus sp. (R. DeBoer, C D A ) . Humboldt Co.: % mile north Kneeland, V I I I 3-61, on Sequoia sempervirens (T. R. Haig, C D A ) . Marin Co.: Blakes Landing, IX-3-61, on Cupressus sp. (U. Nur, C D A ) ; Mill Valley, date ?, on Sequoia sempervirens (G. A. Coleman, C D A ) . Monterey Co.: Carmel, VIII-11—42, on Cupressus macrocarpa (H. H. Keifer, C D A ) ; VII-9-42, on Juniperus sp. (M. R. Bell, C D A ) ; Cypress Point, X - 2 7 - 1 6 , on Cupressus macrocarpa (R. K. Vickery, C D A ) ; Hopkin Marine Laboratory, IX-19-30, on Cupressus macrocarpa (A. C. Browne, C D A ) ; Pacific Grove, X-26-44, on Cupressus macrocarpa (M. R.Bell, C D A ) . Orange Co.: San demente, VII-11-63, on Cupressus macrocarpa (A. Johnson, C D A ) . Riverside Co.: Banning, VIII-26-42, on Juniperus hibernica (C. R. Tower, CDA). San Francisco Co.: San Francisco, VII-2-59, on Cupressus macrocarpa (M. Stufflebeam, C D A ) ; I I 19-29, on Cupressus macrocarpa (H. H. Keifer, C D A ) ; VIII-13-07, on Cupressus macrocarpa (E. M. Ehrhorn, USNM); IX-1-10, on Cupressus macrocarpa (A. Russell, USNM); San Francisco (Golden Gate Park), 1898, on Cupressus macrocarpa (E. M. Ehrhorn, USNM); IX-29-63, on Cupressus sp. (foliage) (D. R. Miller, and J. A. Froebe, U C D ) ; San Francisco (Fort Mason), V I 2-53, on Cupressus sp. (twigs) (R. Bozzini, C D A ) . San Luis Obispo Co.: San Luis Obispo, VI-10-09, on Cupressus macrocarpa (I. J. Condit, USNM); Shell Beach, III—5—52, on Cupressus macrocarpa (W. H. Frey, C D A ) . Santa Clara Co.: Palo Alto, 1911, Cupressus macrocarpa (Childs, U C D ) ; V-24-33, on Cupressus macrocarpa (Burke, U C D ) ; Stanford University, V-16-17, on Sequoia sempervirens (G. F. Ferris, UCD). Santa Cruz Co.: Santa Cruz (nursery), VIII-6-42, on Cupressus macrocarpa (M. R. Bell, C D A ) . Ventura Co.: Camarillo, 11-14-61, on Matthoila sp. (leaves) (W. M. Jones, C D A ) ; Port Hueneme, V-13-63, on Cupressus sp. (G. G. Beevor and B. Osuna, C D A ) ; Ventura, X - 2 3 - 2 9 , on Cupressus sp. (E. L. Smith, C D A ) .

Spilococcus keiferi McKenzie Keifer Mealybug (Figure 179) Spilococcus keiferi McKenzie, 1960. Hilgardia 2 9 ( 1 5 ) : 7 6 1 - 7 6 3 , illus. Type Locality and Host. South Sacramento, Sacramento County, California, on California mugwort, Artemisia vulgaris var. heterophylla (Compositae). North American Distribution. California and Washington. Additional Hosts. Franseria chamissonis (Compositae) and Gramineae (associated with ants). External Features and Habitat. The adult females of this mealybug are covered with a gray, waxy secretion. There are 4 pairs of caudal filaments, all very small, median pair longest; and the legs are reddish. The body contents are green when crushed, this secretion being readily exuded through the ostioles. No ovisac was observed. The species may infest the leaves, stems, or roots of its host. Recognition Characters. Adult females, mounted, 1.90 to 2.90 mm long, 1.00 to 1.60 mm wide; body form broadly oval. Dorsum with number of cerarii reduced, normally 6 to 7 recognizable pairs on abdomen counting forward from anal lobes, absent elsewhere along margin. Anal lobe cerarius with 2 rather small, slender conical setae, 3 to 5 slender auxiliary setae, few scattered trilocular pores; remaining cerarii along abdominal margin with 2 slender setae, these progressively smaller, more slender, usually with 1 or 2 of the pairs more widely separated than in anal lobe pair, no aux-

COLOR PLATE XXIV. The cypress mealybug, Spilococcus implicatus Ferris, on foliage of Monterey cypress, Cupressus macrocarpa (Cupressaceae), San Francisco (Golden Gate Park), San Francisco County, California.

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Fig. 179. Spilococcus keiferi McKenzie, collected on California mugwort, Artemisia vulgaris variety heterophylla (Compositae), south Sacramento, Sacramento County, California.

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iliary setae, scarcely any concentration trilocular pores. Trilocular pores rather evenly beset on dorsum. Minute circular pores scattered. Oral-collar tubular ducts of 2 sizes, larger ones occurring predominantly along body margins, smaller ones rather generally distributed on abdomen, thorax and head. Oral-rim tubular ducts quite numerous, as many as 8 to 10 on each abdominal segment, except the last, scattered over thoracic segments and head. Body setae short and slender. Anal ring apical, of normal form and size for genus; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores numerous, situated predominantly in midregion of abdomen from apical segment forward to fourth, few in lateral areas of last 3 or 4 abdominal segments. Trilocular pores rather numerous, generally distributed over venter, perhaps less abundant in sternal area. Minute circular pores scattered. Oral-collar tubular ducts of 2 sizes, larger ones more numerous, represented in groups in lateral areas of ninth to fourth abdominal segments, on thorax between anterior spiracles and body margin; smaller ones scattered on thorax. Oralrim tubular ducts limited in numbers, situated only along submargin of thorax. Body setae slender, of various lengths, some noticeably longer than those on dorsum. Circulus absent. Legs moderately slender in relation to their length; femora quite stout; hind coxae with cluster of translucent pores; hind tibiae with cluster of translucent pores on distal half of segment; claw without denticle on plantar surface. Antennae 8-segmented, slender. Notes. This species is perhaps most closely related to Spilococcus pressus Ferris. It differs from S. pressus mainly in the possession of an obvious cluster of oralcollar tubular ducts between anterior spiracles and body margin, whereas, in the species last mentioned, this cluster of ducts is absent. California Records (Map 167) Humboldt Co.: 1 mile south Samoa, VIII-12-63, on Franseria chamissonis (D. R. Miller, U C D ) . Los Angeles Co.: Whittier, 11-24-35, on ? (J. D. Maple, U C D ) . Marin Co.: 7 miles east Nicasio, VIII-26-63, on Gramineae (roots—associated with ants) (D. R. Miller, U C D ) . Sacramento Co.: South Sacramento, X-25-45, on Artemisia vulgaris var. heterophylla (H. H. Keifer, CDA).

Spilococcus larreae Ferris Desert Mealybug (Figure 180) Spilococcus larreae Ferris, 1950. Atlas of scale insects of North America, Ser. V:233-234, illus. Type Locality and Host. Whitewater, San Bernardino County, California, on creosote bush, Larrea divaricata (= Larrea tridentata) (Zygophyllaceae).

North American Distribution. Arizona and California. Additional Hosts. Atriplex sp. (Chenopodiaceae) and Prosopis sp.? (Leguminosae). External Features and Habitat. The adult female of this mealybug is finely covered with a smooth, white, waxy bloom. There are 2 small bare areas on dorsum between the head and thorax, and these show as 2 distinct spots. The conspicuous, elongate, slender filaments along body margins are all the same length, and completely encircle the insect. The body color is a brilliant light green, and when crushed the same color fluid is exuded. An ovisac is formed which is 2 - 3 times as long as mature adult female, and this ovisac covers all but the head of the insect. This mealybug occurs on the foliage of its host during the summer months. During the cold winter months it frequents the roots. Eggs and ovisacs are produced in each habitat. In one collection, ants were observed fondling and attending the mealybugs. Recognition Characters. Adult females, mounted, 2.20 to 4.50 mm long, 1.30 to 3.00 mm wide; body form broadly oval. Dorsum with 13-16 pairs of cerarii, some of those along thoracic margins at times so reduced that they are recognizable only with difficulty, although usually present they may disappear entirely. Anal lobe cerarius with 2 quite large conical setae, without accompanying slender auxiliary setae, small cluster trilocular pores, very small area of sclerotization. Anterior to this, cerarii much the same except those along thoracic areas in which setae become smaller and more slender, no auxiliary setae, small cluster trilocular pores; frontal and ocular cerarii normally present, frontal pair each commonly with 2 or 3 conical

TAXONOMY

Fig. 180. Spilococcus (= Larrea tridentata) California.

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larreae Ferris, collected on creosote bush, Larrea divaricata (Zygophyllaceae), at Whitewater, San Bernardino County,

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setae, ocular pair each usually with 2 conical setae, seldom with 3, these nearly as large as those of anal lobe cerarii, no auxiliary setae, small cluster trilocular pores. Trilocular pores evenly distributed over dorsum. Minute circular pores numerous, scattered over dorsal surface. Oral-collar tubular ducts absent except, perhaps, for a very few along margin of last 2 abdominal segments. Oral-rim tubular ducts comparatively small, extremely numerous, distributed in bands and groups over entire dorsum. Body setae small and slender. Anal ring situated close to posterior apex of abdomen, quite large, sclerotized band rather narrow, with few pores; each of its 6 setae slightly longer than greatest diameter of ring. Venter with multilocular disk pores abundant, situated in midregion of abdomen from apical forward to circulus, not occurring in lateral areas. Trilocular pores scattered over entire venter. Minute circular pores scattered over venter. Oral-collar tubular ducts situated in midregion and lateral areas of most abdominal segments, absent elsewhere on venter. Oral-rim tubular ducts of same size as those on dorsum, occur in considerable numbers in lateral areas of abdomen as far forward as prothorax, in small numbers in sternal areas of thoracic segments, and on head. Body setae slender, of various lengths, some noticeably longer than those on dorsum. Circulus comparatively small, appearing capable of folding along intersegmental line between which it lies. Legs comparatively large; hind coxae without translucent pores; hind tibia with small translucent pores near distal end of segment; claw without denticle on plantar surface. Antennae 8-segmented, rather slender. Notes. This species is related to Spilococcus steelii (Cockerell and Townsend), but differs from it in having the frontal cerarius definitely developed, with 2 or 3 conical setae set close together, whereas, in the species last mentioned, the frontal cerarius is indefinite, represented by a scattered group of 3 or more widely separated conical setae. In addition, S. larreae possesses dorsal oral-rim tubular ducts on prothorax and head, whereas in S. steelii these structures are absent in these areas. California Records (Map 168) Imperial Co.: Holtville, X-10-61, on Larrea divericata (C. R. Waegner, CDA); 111-17-64, on Larrea tridentata (R. A. Flock, CDA); 8 miles east Holtville, IV-8-63, on Prosopis sp. ? (R. L. Westcott, UCD); Saltan City, 111-27-64, on Larrea sp. (G. G. Beevor, CDA); 2 miles east Plaster City, VI-21-64, on Atriplex canescens and Larrea sp. (D. R. Miller, UCD). Inyo Co.: Death Valley, date ?, on Larrea sp. (G. F. Ferris, UCD). Kern Co.: Mojave near Red Rock Canyon, X-1739, on Larrea tridentata (J. W. Dixon, CDA). Riverside Co.: Desert Center (Gruendyke's Well), VI-1918, on Larrea sp. (G. F. Ferris, UCD); Mecca, VII-27-62, on Larrea sp. (K. Hoffmaster, CDA);

White Water, IV-14-35, on Larrea divaricata (J. D. Maple, UCD); X-l-55, on Larrea sp. (W. D. Dyer, LEM). San Bernardino Co.: 26 miles southwest Baker, VI-11-63, on Larrea divaricata (foliage) (D. R. Miller, UCD); near Victorville (Apple Valley), V16-48, on Larrea sp. (W. D. Dyer, CDA); San Diego Co.: Borrego Springs, XI-24-58, on Larrea tridentata (R. J. Buckner and M. Johnson, CDA) ; 7 miles southeast Borrego Springs, 1-27-65, on Larrea tridentata (crown and roots) (D. R. Miller, UCD).

Spilococcus parkeri McKenzie Parker Mealybug (Figure 181) Spilococcus parkeri McKenzie, 1962. Hilgardia 32(14):682-683, 685, illus. Spilococcus parkeri McKenzie, 1964. Ibid. 35(10): 269. Type Locality and Host. Pioche, Lincoln County, Nevada, on Gutierrezia microcephala (Compositae). North American Distribution. California and Nevada. Additional Hosts. Gutierrezia californica ? (Compositae) and Stillingia linearifolia (Euphorbiaceae). External Features and Habitat. This mealybug is covered with a smooth, white, waxy secretion; 2 or 3 pairs of rather small, inconspicuous caudal filaments; body color red; body contents when crushed reddish; and legs red. Cast skins are often found under rocks

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Fig. 181. Spilococcus parked McKenzie, collected on Gutierrezia californica? (Compositae), northeast Gilman Hot Springs, Riverside County, California. (Illustration prepared from type material.)

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and debris at a maximum distance of 1 foot away from the host. An ovisac is formed. The species is found primarily on the crown, secondarily on the roots of its host. An ant, Solenopsis xyloni, was found in association with this mealybug. Recognition Characters. Adult females, mounted, 1.40 to 3.60 mm long, 0.90 to 2.40 mm wide; body form broadly oval. Dorsum with number of cerarii reduced, normally 7 or 8 pairs on abdomen counting forward from anal lobes, possibly 4 or 5 on thorax and head, these difficult to discern because cerarian conical setae usually much more slender and generally situated farther apart, total recognizable cerarii running as high as possibly 13 or 14 pairs. Anal lobe cerarius with 2 or 3 rather slender conical setae, 2 or 3 more slender auxiliary setae, rather concentrated group trilocular pores, no sclerotization; remaining cerarii with usually 2 slender conical setae, these progressively smaller, no auxiliary setae, scarcely any concentration trilocular pores. Trilocular pores evenly distributed on dorsum. Usually 2 or 3 dorsal, submarginal multilocular disk pores situated on sixth and eighth abdominal segments. Minute, circular pores scattered over dorsal surface. Oral-collar tubular ducts absent. Oral-rim tubular ducts rather numerous, 13 to 20 observed, average 16.1, on fourth, fifth, sixth, and eighth abdominal segments, 7 or 8 on seventh, absent on ninth, scattered over thoracic segments and head. Body setae small and slender. Anal ring apical, of normal form and size for genus; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores fairly numerous, extending across abdominal segments from apical forward to fifth segments, few (3 or 4 pores only) on fourth segment. Trilocular pores rather sparsely distributed, especially in sternal area. Minute, circular pores generally scattered on venter. Oralcollar tubular ducts considerably numerous, especially along posterior margin of ninth forward to third abdominal segments, few scattered on thorax, absent on head. Oral-rim tubular ducts same size as those on dorsum occur in small numbers in lateral areas of eighth forward to sixth segments, more numerous on anterior abdominal segments and thorax, few scattered in sternal area and on head. Body setae slender, of various lengths, some noticeably longer than those on dorsum. Circulus comparatively large, transversely-oval, situated on fourth abdominal segment, usually not divided by segmental line across which it lies. Legs well developed, comparatively long; hind coxae with cluster of translucent pores at base; hind tibiae with irregular cluster or 2 of translucent pores situated on apical half of segment; slight denticular swelling on plantar surface of claw, or this completely absent. Antennae 8-segmented. Notes. This species appears quite closely related to Spilococcus parvicirculus McKenzie, but differs mainly in possessing more numerous dorsal oral-rim

tubular ducts (13 to 20, average 16.1) on each of the fourth, fifth, sixth, and eighth abdominal segments; by having at least a few marginal cerarii on head and thorax; and usually a larger circulus. Spilococcus parvicirculus, on the other hand, usually possesses not more than 6 to 10, average 8.3, dorsal oral-rim tubulars on each of the fourth, fifth, sixth, and eighth abdominal segments; no marginal cerarii on head and thorax; and usually a very small, transversely-oval, circulus. Spilococcus parkeri, originally described from Nevada, has been subsequently found in California. California Records (Map 169) Riverside Co.: 3.4 miles northeast Gilman Hot Springs, III-9-63, on Guiterrezia californica (?)10 (D. R. Miller, U C D ) .

Spilococcus parvicirculus McKenzie Small-Circulus Mealybug (Figure 182) Spilococcus parvicirculus McKenzie, 1960. Hilgardia 2 9 ( 1 5 ) : 7 6 2 - 7 6 4 , illus. Type Locality and Host. Ontario, San Bernardino County, California, on Franseria acanthicarpa (Compositae). North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. No available infor10 May be Stillingia linearifolia (roots) (Euphorbiaceae). This plant was taken in recollection. Identified by June McCaskill.

TAXONOMY

Fig. 182. Spilococcus parvicirculus McKenzie, collected on Franseria (Compositae), at Ontario, San Bernardino County, California.

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acanthicarpa

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mation on external appearance of this mealybug. The species presumably occurs above ground on its host. Recognition Characters. Single available adult female, mounted, 2.25 mm long. 1.30 mm wide; body form broadly oval. Dorsum with cerarii rather poorly developed, 7 to 8 recognizable pairs on abdomen counting forward from anal lobes, absent on thorax and head. Anal lobe cerarius with 2 rather slender conical setae, 2 or 3 more slender auxiliary setae, few scattered trilocular pores; remaining cerarii along abdominal margin with 2 slender conical setae, these progressively smaller away from anal lobes, no auxiliary setae, scarcely any concentration trilocular pores. Trilocular pores evenly beset over entire dorsum. Oral-collar tubular ducts of 2 sizes, very few, situated predominantly along margin of last 2 or 3 abdominal segments, absent elsewhere on dorsum. Oral- rim tubular ducts quite numerous, as many as 4 to 10 on each abdominal segment, except last, scattered over thoracic segments and head. Body setae small and slender. Anal ring apical, normal in form and size for genus; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores fairly numerous in midregion of abdomen from apical to posterior margin of fifth abdominal segment. Trilocular pores rather evenly beset on venter except for certain "clear" areas on thorax where they are absent. Oralcollar tubular ducts of 2 sizes situated from apical forward to fourth abdominal segments, across segments and in clusters along margin, larger ones more numerous. Oral-rim tubular ducts of same size as those on dorsum, occur in small numbers in lateral areas of most segments from eighth abdominal forward to head, scattered in sternal area, and about spiracles. Body setae slender, many noticeably longer than those on dorsum. Circulus very small, transversely oval (from whence the specific name was derived). Legs well developed, moderately slender in relation to their length; hind coxae with cluster translucent pores at base; hind tibiae without translucent pores; claw without denticle on plantar surface. Antennae 8-segmented, slender, second and third segments from base longest. Notes. This species appears to be quite closely related to Spilococcus parkeri McKenzie, but differs from it mainly in having fewer dorsel oral-rim tubular ducts (6 to 10, average 8.3) on each of the fourth, fifth, sixth, and eighth abdominal segments; and by the absence of marginal cerarii on the thorax and head. Spilococcus parkeri, on the other hand, possesses numerous dorsal oral-rim tubulars (13 to 20, average 16.1) on the abdominal segments previously mentioned, as well as at least a few marginal cerarii on the thorax and head. California Records (Map 170) San Beranardino Co.: Ontario, VI-22-59, on Franseria acanthicarpa (G. M. Harper, C D A ) .

Map 170. Distribution in California of Spilococcus parvicirculus McKenzie.

Spilococcus pressus Ferris Pressus Mealybug (Figure 183) Spilococcus pressus Ferris, 1950. Atlas of scale insects of North America, Ser. V: 23 5 - 2 3 6, illus. Type Locality and Host. Lake on Big Creek, Santa Cruz County, California, on wood violet, Viola sarmentosa (Violaceae). North American Distribution. Known only from California. Additional Hosts. Achillea millefolium (Compositae), Althaea rosea (Malvaceae), Aster chilensis and A. spinosus (Compositae), Crassula sp. (Crassulaceae), Eremocarpus setigerus (Euphorbiaceae), Gossypium sp. (Malvaceae), Nerium oleander (Apocynaceae), Oenothera sp. (Onagraceae), Phacelia sp. (Hydrophyllaceae), Sida sp. (Malvaceae), Stachys bullata (Labiatae), Viola sarmentosa (Violaceae), and "turkey weed" (specific plant identification unknown). External Features and Habitat. The adult female of this species is lightly dusted with a smooth, gray, waxy secretion. The body color is greenish; body contents when crushed green; body shape oval; legs white. There are 3 pairs of short, thin, caudal filaments—the posteriormost pair slightly longer than the other 2. A filamentous ovisac is produced which is one-third the length of mature adult female and covers only posterior part of abdomen. This mealybug occurs on the crown of its host. Ferris (1950) indicates that dates of collection range from early June to September. Recognition Characters. Adult females, mounted, 1.95 to 2.50 mm long, 1.10 to 1.40 mm wide; body form broadly oval.

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Fig. 183. Spilococcus pressus Ferris, collected on wood violet, Viola sarmentosa (Violaceae), at lake on Big Creek, Santa Cruz County, California.

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Dorsum with 8 - 9 recognizable cerarii on abdomen and metathorax, 0 - 3 cerarii anterior to metathorax, ocular cerarius generally present, frontal cerarius absent. Anal lobe cerarius with 2 quite small, slender conical setae, 1 - 3 slender auxiliary setae, very few scattered trilocular pores; remaining cerarii with 2 very small conical setae, no auxiliary setae, very few dispersed trilocular pores. Trilocular pores sparsely scattered over dorsum. Oral-collar tubular ducts absent. Oral-rim tubular ducts rather few, somewhat variable in number and position, about 7 - 8 across any one abdominal segment, except the last, lightly scattered in thoracic and head regions. Body setae relatively few, small, and slender. Anal ring apical, normal for genus; each of its 6 setae only slightly longer than greatest diameter of ring. Venter with multilocular disk pores relatively few, there being scarcely more than 30 on any one abdominal segment, extending forward from apical segment in very small numbers to fifth or perhaps fourth segment, absent elsewhere on body. Trilocular pores sparsely distributed, certain "clear" areas evident especially on thorax. Oral-collar tubular ducts situated in small numbers in midregion of abdomen, in small clusters of perhaps not more than 10 in lateral areas of segments 8-4, occasional duct occurs in lateral areas of thorax, absent on head. Oral-rim tubular ducts very few, usually not more than 1 in lateral areas of 2 or 3 abdominal segments, 1 - 3 in areas opposite each spiracle on thorax. Body setae for most part short and slender, certain ones appearing definitely longer than those on dorsum. Anal lobes with very small and indistinct ventral sclerotization at base of one of minor setae. Circulus absent. Legs well developed; hind coxae with cluster of pores at base; hind tibiae without translucent pores; claw without denticle on plantar surface. Antennae normally 8-segmented. Notes. This species is related to Spilococcus keiferi McKenzie, but differs mainly in the absence of a ventral cluster of oral-collar tubular ducts between anterior spiracles and body margin, whereas in the species last mentioned, this pore cluster is quite obvious. California Records (Map 171) Contra Costa Co.: Antioch, IX-1936, on Oenothera sp. (H. M. Armitage, C D A ) . Imperial Co.: Brawley, 1-6-64, on Althaea sp. (R. A. Flock and J. Carr, C D A ) ; X-16-64, on Achillea millefolium (Mohammed and Locker, C D A ) ; X - 2 2 - 6 4 , on Nerium oleander (R. A. Flock and J. V. Taylor, C D A ) ; XI-10-64, on Crassula sp. (T. B. Gallion and Heinz, C D A ) ; Calexico, X I - 1 8 64, on Aster chilensis (R. A. Flock, C D A ) ; X I - 1 8 64, on Gossypium sp. (R. A. Flock and Cano, C D A ) ; Callpatria, XI-24 and 25-64, on Althaea rosea ( R . A. Flock, C D A ) ; El Centro, III-6-66, on Aster spinosus (J. V. Taylor and C. R. Waegner, C D A ) ; Imperial, V-18-66, on Nerium oleander (J. L. Johnson and E. Menvielle, C D A ) ; and Winterhaven, XI-1-65, on Gossypium sp. (T. Plumb, C D A ) .

Orange Co.: 7 miles east San Juan Capistrano, VI-22-64, on Eremocarpus setigerus (crown, foliage, and associated with ant, Crematogaster coarctata vermiculata) (D. R. Miller, U C D ) . Riverside Co.: Riverside, Citrus Experiment Station Grounds, VIII-19-24, on Eremocarpus setigerus (J. C. Chamberlin, U C D ) . San Bernardino Co.: 1 mile west Joshua Tree, IV9 - 6 3 , on Phacelia sp. (D. R. Miller, U C D ) ; Upland, VIII-19-25, on "turkey weed" (C. P. Clausen, CDA). San Diego Co.: 5 miles north Escondido, 111-2764, on Sida sp. (D. R. Miller and J. A. Froebe, UCD). Santa Clara Co.: Stanford University (Corte Madera Creek), VII-21-17, on Stachys bullata (G. F. Ferris, U C D ) . Santa Cruz Co.: Lake on Big Creek, VI-10-17, on Viola sarmentosa (G. F. Ferris, U C D ) .

Spilococcus prosopidis (Cockerell) Mesquite Mealybug (Figure 184) Dactylopius prosopidis Cockerell, 1896. Canad. Ent. 28:224. Pseudococcus prosopidis (Cockerell), Fernald, 1903. Mass. Agr. Expt. Sta. Spec. Bull. 88, p. 108. Spilococcus prosopidis (Cockerell), Ferris, 1950. Atlas of scale insects of North America, Ser. V : 2 3 7 - 2 3 8 , illus. Type Locality and Host. Mesilla, Dona Ana County, New Mexico, on Prosopis sp. (Leguminosae).

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Fig. 184. Spilococcus prosopidis (Cockerell), collected on Prosopis sp. (Leguminosae), ^ at Andrade, Imperial County, California. (Illustration prepared from specimens col- ' lected on Prosopis sp. [Leguminosae], at El Paso, El Paso County, Texas.)

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North American Distribution. Arizona, California, New Mexico, and Texas. Additional Hosts. Phoradendron sp. (Loranthaceae). External Features and Habitat. According to Cockerell (1896), "the female forms a dense cushion of white cottony matter, on which it rests; this cushion is visible all around the margin of the insect, and does not at first protrude greatly behind. In it are laid the very pale greenish-yellow eggs. Eventually the females become dark slate-gray, and have a cross of white secretion on the hind end. They at this period possess a Pulvinaria-like ovisac . . . Boiled in caustic soda, they stain the fluid cochineal-red, and turn orange." The insect infests the twigs of its host, and, according to the original description, are attended by ants. Recognition Characters. Adult females, mounted, 2.00 to 2.50 mm long, 1.60 to 2.00 mm wide; body form broadly oval. Dorsum with recognizable cerarii much reduced, scarcely more than 7 pairs distinguished counting forward from anal lobes, this due to fact setae become widely separated and very small, and finally merge with body setae. Anal lobe cerarius with 2 conical setae, 3-4 slender auxiliary setae, very few scattered trilocular pores; remaining cerarii with 2 slender conical setae, no auxiliary setae, little or no concentration trilocular pores. Trilocular pores so arranged they are practically absent except in 2 longitudinal bands on each side; evidently these bare areas correspond to areas which lack waxy secretion in living insects. Minute circular pores scattered. Oral-collar tubular ducts absent. Oralrim tubular ducts in marginal row from eighth abdominal segment forward to head, absent elsewhere on dorsum. Body setae relatively few, all quite small, with distribution similar to trilocular pores. Anal ring apical, normal for genus; each of its 6 setae slightly longer than greatest diameter of ring. Venter with multilocular disk pores present in considerable numbers in midregion of abdomen from apical forward to posterior margin of third segment, occasionally in small numbers in lateral areas of some of the abdominal segments, absent elsewhere on venter. Trilocular pores moderately distributed, certain "clear" areas evident, especially on thorax where these pores are absent. Minute circular pores scattred. Oral-collar tubular ducts very small, occur in few numbers in midregion of abdomen, in clusters near lateral margins on all segments as far forward as mesothorax. Oral-rim tubular ducts, similar in size to those on dorsum, occur in variable numbers in lateral areas of all abdominal segments, except the last, forward to prothorax and head, scattered in sternal areas of thoracic segments. Body setae short and slender, some slightly longer than those on dorsum. Anal lobes with very small, sclerotized bar which extends in at base of one of minor setae. Circulus broadly oval, capable of folding along

intersegmental line. Legs comparatively short and stout; hind coxa with few translucent pores at base; hind tibia with similar pores near distal end of segment. Claw without denticle on plantar surface. Antennae normally 8-segmented. Notes. This species seems rather closely related to Spilococcus andersoni (Coleman), but differs in having not more than 6 or 7 definite pairs of cerarii, these confined to the abdominal segments, whereas in the species last mentioned there are about 15 pairs of definite cerarii, many of which occur on thorax and head. In addition, S. prosopidis occurs only on Prosopis spp., while S. andersoni is found only on members of the Cupressaceae, Taxaceae, and Taxodiaceae. California Records (Map 172) Imperial Co.: Andrade, 111-30-45, on Prosopis sp. (C. G. Anderson and A. J. Hanson, CDA). San Bernardino Co.: Kingston, X-24-59, on Prosopis sp. (G. M. Harper, CDA). San Diego Co.: Jacumba, XII-4-51, on Phoradendron sp. (fruit cluster) (G. W. Schwegel, CDA).

Map 172. Distribution in California of Spilococcus prosopidis (Cockerell).

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Spilococcus quercinus McKenzie Chaparral Mealybug (Figure 185) Spilococcus quercinus McKenzie, 1962. Hilgardia 32(14):684-686, illus. Type Locality and Host. Big Rock Creek, San Gabriel Mountains, Los Angeles County, California, on oak, Quercus sp. (Fagaceae). North American Distribution. Known only from California.

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Fig. 185. Spilococcus quercinus McKenzie, collected on oak, Quercus sp. (Fagaceae), at Big Rock Creek, San Gabriel Mountains, Los Angeles County, California.

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Additional Hosts. Ceanothus cuneatus (Rhamnaceae). External Features and Habitat. No available information on external appearance of this species. It is suspected that the mealybug feeds on stems and foliage of its host. Recognition Characters. Adult females, mounted, 2.00 to 4.20 mm long, 1.10 to 2.10 mm wide; body form broadly oval. Dorsum with cerarii rather poorly developed, variable in number, usually 7-8 pairs on abdomen, 0 - 2 on thorax, 0 - 1 pair on head (usually preocular), these difficult to discern because cerarian conical setae usually more slender, often situated farther apart, total recognizable cerarii ranging from 7 to 11. Anal lobe cerarius with 2 rather small, slender conical setae, 4 or 5 slender auxiliary setae, few scattered trilocular pores; remaining cerarii along body margin with but 2 conical setae, these more slender and at times, more widely separated than anal pair, strongly resembling body setae, difficult to recognize, no auxiliary setae, scarcely any concentration trilocular pores. Trilocular pores evenly distributed on dorsum. Minute circular pores lightly scattered. Oral-collar tubular ducts absent. Oral-rim tubular ducts numerous, arranged in broken double row of from 30 to 38, or perhaps even more, across each abdominal segment, except the last (0-4 ducts), fairly numerous on thoracic segments, few scattered on head. Body setae small and slender. Anal ring apical, of normal form and size for genus; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores considerably numerous, situated in midregion of abdomen from apical forward to third segment (usually only 1 or 2 pores present on this segment), absent on thorax and head. Trilocular pores evenly distributed on venter, except for certain areas of thorax and head, where they appear more sparse. Minute circular pores lightly scattered. Oral-collar tubular ducts occur on abdomen in same general area as multilocular disk pores, except on second and third segments, considerably numerous in clusters of lateral areas of ninth to fifth segments, apparently absent on thorax and head. Oral-rim tubular ducts of same size as those on dorsum occur in small numbers submarginally on abdomen, thorax and head, and in area about spiracles. Body setae evenly distributed on venter, except in certain areas of thorax and head, where they appear more sparse. Circulus normally appearing broadly oval, capable of folding along intersegmental line of segments across which it lies. Legs moderately slender in relation to length; hind coxa with cluster of translucent pores at base; hind tibia with translucent pores, irregularly arranged on distal half of segment; claw with very small denticle on plantar surface. Antennae normally 8-segmented. Notes. This species is perhaps most closely related to Spilococcus cactearum McKenzie, but differs from

it chiefly in the greater number of dorsal oral-rim tubular ducts (20 or more) on abdomal segments; by the presence of ventral multilocular disk pores on the fourth abdominal segment; and translucent pores on hind tibiae more generally scattered over entire segment. Spilococcus cactearum, on the other hand, has fewer dorsal oral-rim tubulars (less than 10) on abdominal segments; lacks ventral multilocular disk pores on fourth abdominal segment; and with a small irregular cluster of translucent pores on apical half of hind tibiae. The hosts of these 2 mealybugs are also quite different, S. cactearum infesting cactus, while S. quercinus has been collected on oak and Ceanothus. California Records (Map 173) Los Angeles Co.: Big Rock Creek, San Gabriel Mountains, IV-1926, on Quercus sp. (J. C. Chamberlain, C D A ) . Ventura Co. : Santa Paula, date ? , on Ceanothus cuneatus (E. O. Essig, CAS).

Spilococcus sequoiae (Coleman) Redwood Mealybug (Figure 186) Dactylopius sequoiae Coleman, 1901. Proc. Cailf. Acad. Sci., (Ser 3, Zool.) 2 ( 1 1 ) : 409-420, illus. (in part, as to the female). Pseudococcus sequoiae (Coleman), Fernald, 1903. Mass. Agr. Expt. Sta. Spec. Bull. 88, p. 109. Pseudococcus sequoiae (Coleman), Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., pp. 53-54, illus. Spilococcus sequoiae (Coleman), Ferris, 1950. Atlas

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Fig. 186. Spilococcus sequoiae (Coleman), collected on redwood, Sequoia sempervirens (Taxodiaceae), at Stanford University, Santa Clara County, California (Topotype).

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of scale insects of North America, Ser. V.239240, illus. Type Locality and Host. Stanford University, Santa Clara County, California, on redwood, Sequoia sempervirens (Taxodiaceae). North American Distribution. California and Mexico. Additional Hosts. Cupressus macrocarpa and Cupressus spp. (Cupressaceae). External Features and Habitat. A long, slender, whitish, firm ovisac is formed at maturity. The species occurs on the foliage of its host. Recognition Characters. Adult females, at maturity, 2.25 to 3.40 mm long, 1.10 to 2.20 mm wide; body form broadly oval. Dorsum with cerarii very weakly developed, 3, or perhaps at times 4, pairs on abdomen, on head region, counting from frontal cerarius, 4 pairs often recognizable, total of scarcely more than 8 pairs. Anal lobe cerarius with 2 quite small, conical setae, 3-4 slender auxiliary setae, very few scattered trilocular pores; remaining cerarii on abdomen, prothorax, and head each with 2 very small conical setae, no slender auxiliary setae, 3-4 trilocular pores. Trilocular pores very sparse, though evenly distributed on dorsum. Few multilocular disk pores come over to dorsum from ventral clusters on seventh, eighth, and ninth abdominal segments. Oralcollar tubular ducts absent. Oral-rim tubular ducts either entirely absent or only 1 or 2 situated on head. Body setae relatively few, small, and slender. Anal ring apical, normal in size and cellularity for genus; each of its 6 setae twice as long as greatest diameter of ring. Venter with multilocular disk pores in small numbers in midregion of abdomen from posterior apical segment forward to fifth segment, in noticeable clusters in lateral areas of each abdominal segment, thoracic region forward to mesothorax. Trilocular pores sparsely distributed over entire venter. Oral-collar tubular ducts situated in small numbers in midregion of abdominal segments forward to sixth segment and amongst clusters of lateral multilocular disk pores. Oral-rim tubular ducts in clusters mesad of marginal clusters of multilocular disk pores on abdominal segments and in area about spiracles on thoracic segments. Body setae for most part short and slender, certain ones appearing slightly longer than those on dorsum. No sclerotized area present on anal lobes. Circulus absent. Legs well developed; hind coxae with numerous translucent pores at base, fore- and mid-coxae with few, inconspicuous translucent pores; hind tibiae with few inconspicuous, translucent pores scattered along entire segment, fore- and mid-tibiae with slight indication translucent pores; claw without denticle on plantar surface. Antennae normally 8segmented. Notes. According to Ferris (1950), the describer of this species included 3 species in his type material. The male ascribed to this species was that of Puto cupressi (Coleman). The females involved 2 species on the same slides. A type selection was made by

Ferris from these slides which fixed the name S. sequoiae. The other species was Spilococcus implicatus, described by Ferris in 1950. The combination of lateral clusters of multilocular disk pores on venter of abdomen and thorax, together with the absence of a circulus, is quite distinctive of this species. This mealybug may occur in company with S. implicatus, both maturing at the same time and apparently quite indistinguisable from each other in life. California Records (Map 174) Alameda Co.: Berkeley, date ?, on Cupressus macrocarpa (E. O. Essig, CAS). Marin Co.: Blake's Landing, IX-3-61, on Cupressus sp. (U. Nur, UCD). Monterey Co.: Pacific Grove, VII-23-62, on Cupressus macrocarpa (V. B. Whitehead, UCD); 6 miles south Big Sur, VIII-10-62, on Sequoia sempervirens (E. I. Schlinger, UCD). San Francisco Co.: San Francisco, IX-9-37, on Cupressus macrocarpa (E. S. Ross, CAS). Santa Clara Co. : Stanford University, IV-20-1900, on Sequoia sempervirens (G. A. Coleman, UCD); V-14—45, on Sequoia sempervirens (G. F. Ferris, UCD); IX-17-16, on Sequoia sempervirens (G. F. Ferris, UCD).

Spilococcus ventralis McKenzie, new species Burro-weed Mealybug (Figure 187) Type Locality and Host. Type and paratype adult females on burro-weed, Franseria dumosa (Compositae), 6 miles north Inyokern, Kern County, Cali-

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Fig. 187. Spilococcus ventralis McKenzie, new species, collected on burro-weed, Franseria dumosa (Compositae), north of Inyokern, Kern County, California.

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fornia, collected December 28, 1964, by D. R. and J. F. Miller. These are the only known specimens of the species. Type Material. Holotype adult female (single specimen on slide) and female paratypes in museum, University of California, Davis. Paratypes in United States National Collection of Coccoidea (U. S. National Museum), Washington, D. C., and in collection of California State Department of Agriculture, Sacramento. North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. This mealybug is lightly covered with a smooth, white, powdery secretion. There are 2 faint spots on dorsum of metathorax. The body form is subcircular; body color red; body contents orange when crushed; legs reddish. There are 2 pairs of thin, minute, caudal filaments. A filamentous ovisac is produced by adult female, and a male sac is formed. The mealybug occurs on the crown and roots of host, and sometimes in tunnels of wood-boring beetles. Ants were found in association with this collection. Recognition Characters. Adult females quite small for members of this group, mounted, 1.80 to 1.95 mm long, 1.10 to 1.25 mm wide; body form broadly oval. Dorsum with cerarii somewhat variable in number, not more than 12 pairs, including 8 abdominal pairs counting forward from anal lobes, 3 - 4 thoracic pairs, these often represented by a single conical seta. Anal lobe cerarius with 3 - 5 moderately elongate conical setae, several longer, more slender auxiliary setae, few obviously associated trilocular and minute circular pores. Remaining abdominal cerarii each with 2 or 3 conical setae, progressively slightly smaller with distance from anal lobes, no auxiliary setae, slight concentration trilocular pores; cerarii on thorax similarly constructed, often represented by a single seta. Dorsum rather evenly beset with trilocular pores; less numerous minute circular pores. Oral-rim tubular ducts absent on dorsal surface. A few oral-collar tubular ducts evident along margin of last 4 or 5 abdominal segments, absent elsewhere on dorsum. Body setae short and slender. Anal ring apical, normal for genus; each of its 6 setae about twice as long as diameter of ring. Venter with multilocular disk pores present in considerable numbers extending across abdominal segments from apical to posterior margin of fourth segment, 1 or 2 pores on third segment, few pores in sternal area of metathorax, absent elsewhere on venter. Trilocular pores evenly beset on venter, except for certain "clear" areas on thorax and head. Minute circular pores sparsely distributed. Oral-rim tubular ducts occur in few numbers in lateral areas of abdo-

men, especially abundant between hind and mid-pair of legs, and mid-pair of legs and anterior spiracle, few along submargin of prothorax, absent on head. Oral-collar tubular ducts fairly numerous on abdomen, scattered on thorax, absent on head. Most body setae slightly longer than those on dorsum. Circulus moderately large, oval, faintly divided by intersegmental line. Legs well developed; hind coxa with numerous translucent pores at base; distal half of hind tibia with 1 or 2 irregular clusters of similar pores; denticular swelling absent on plantar surface of claw. Antennae 8-segmented. Notes. This species appears rather closely related to Spllococcus townsendi (Cockerell), but differs in the absence of obvious clusters of small oral-collar tubular ducts in ventrolateral areas of thorax; 8-segmented antenna; anal lobe cerarius with 3 - 5 moderately elongate conical setae, several longer, more slender auxiliary setae; and known only from Franseria dumosa (Compositae) in California. Spilococcus townsendi, on the other hand, possesses obvious clusters of small oral-collar tubular ducts in ventrolateral areas of thorax; possesses a 7-segmented antenna; anal lobe cerarius with only 2 moderately stout, conical setae, 1 or more slender auxiliary setae; as far as known occurs only on Fouquieria splendens (Fouquieriaceae) in southwest United States. California Records (Map 175) Kern Co.: Inyokern, XII-28-64, on Franseria dumosa (D. R. and J. F. Miller, U C D ) .

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STEMMATOMERINX Stemmatomerinx Ferris, 1950. Atlas of the scale insects of North America, Ser. V:245. Genotype. Stemmatomerinx decorata Ferris. Recognition Characters. Pseudococcidae with dorsal ostioles present, the posterior pair not too well developed. Differing from other members of family in having cerarian setae and some dorsal body setae, enlarged, stout, in form of a truncate cone, the base of each of these modified setae surrounded by a ring bearing from 3 - 7 small pores. Dorsal cerarii difficult to determine in type species, since groups are in part somewhat diffuse, but there seems to be 17 pairs. Tubular ducts of oral-collar type present. Multilocular disk pores, and quinquelocular pores present. Tubular ducts of oral-collar type present. Circulus present in type species. Legs well developed; claw with denticle on plantar surface. Notes. Pseudococcidae relating to the so-called "Phenacoccus" series, that is, with a denticle on plantar surface of claw, and 9-segmented antennae. According to Ferris (1950), "This genus is evidently somewhat like Synacanthococcus, known only from the Philippine Islands, but differs from the latter in having the enlarged dorsal and cerarian setae truncate, and surrounded basally by a ring of small pores. Otherwise there seems to be nothing else known that is at all similar."

Stemmatomerinx decorata Ferris Truncate Seta Mealybug (Figure 188) Stemmatomerinx decorata Ferris, 1950. Atlas of scale insects of North America, Ser. V : 2 4 6 247, illus. Type Locality and Host. One mile or less north of Beatty, Nye County, Nevada, on Distichlis sp. (Gramineae). North American Distribution. California, Nevada, New Mexico, and Texas. Additional Hosts. Undetermined grass (Gramineae). External Features and Habitat. According to Ferris (1950), the adult females of this species are slender, firmly felted, and beset with long, slender, stiff threads of wax, especially at the posterior end. It is reported that the insect is almost entirely enclosed at the anterior end of the sac, but that it may fall out of this sac very easily when disturbed. This mealybug occurs on the foliage of its host, usually on the upper side of the leaves. Recognition Characters. Adult females, mounted,

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3.00 to 4.00 mm long, 1.60 to 1.70 mm wide; body form tending to be somewhat slender in fully expanded specimens. Dorsum with number cerarii difficult to determine, since groups, in part, somewhat diffuse, total appears to be 17. Anal lobe cerarius with 3 large, truncate setae set in circular, sclerotized area that bears at most but very few trilocular pores. Remaining cerarii with 2 - 3 modified setae, groups on abdominal segments and in head region surrounded by indistinct, slightly sclerotized area. Trilocular pores sparsely beset on dorsum. Multilocular disk pores few, very inconspicuous, situated on posterior abdominal segments. Tubular ducts of 2 sizes, large ones with slight oral collar occur in considerable numbers over entire dorsum, except posterior apical abdominal segment. Oral-rim tubular ducts absent. Enlarged truncate setae of dorsum, each with wreath about base bearing 3-7 small pores, most commonly 5, these pores possibly modified trilocular pores, arranged in 4 rows, each row, anterior to ninth abdominal segment, with single seta on each body segment, except in head region where rows become irregular. On seventh and eighth abdominal segments enlarged setae of 2 submedian rows set close together in pairs on small sclerotized areas. Few extremely small, conical setae scattered over dorsum, and an occasional larger, conical seta, especially near cerarii. Anal ring somewhat elongate; each of its 6 setae but slightly longer than diameter of ring itself. Venter with multilocular disk pores rather small, numerous, situated both medially and in lateral areas of abdominal segments posterior to circulus. Quinquelocular pores small, inconspicuous, in midregion of body from sixth abdominal segment forward to head. Trilocular pores very few, sparsely situated along submargin of body. Oral-collar type ducts of smaller size situated in small numbers over most of venter; larger-sized ones occur along lateral margins from about fifth abdominal segment forward to head. Body setae slender and various lengths. Circulus rather small, oval. Legs well developed; claw with distinct but small denticle on plantar surface. Antennae 9-segmented, slender. Notes. This species is markedly different from any other California pseudococcid in having the cerarian setae and some of the dorsal body setae enlarged, stout, and in the form of a truncate cone, the base of each of these modified setae being surrounded by a ring bearing 3 - 7 small pores. California Records (Map 176) Mono Co.: Laws, VII-27-47, on Distichlis sp. (G. F. Ferris, U C D ) .

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Fig. 188. Stemmatomerinx decorata Ferris, collected on salt grass, Distichlis sp." (Gramineae), at Laws, Mono County, California.

TRIDISCUS Tridiscus Ferris, 1950. Atlas of the scale insects of North America, Ser. V:248. Tridiscus Ferris, 1953. Ibid., Vol. VI:477. Genotype. Trionymus distichlii Ferris. Recognition Characters. Pseudococcidae with ostioles either completely absent or reduced, anterior pair apparently absent, posterior pair very small and detectible only in favorable specimens. Dorsum with cerarii present only on anal lobes, here represented by slender, paired setae. All pores and ducts—trilocular, multilocular, and oral-collar tubulars—of normal form. Anal ring well developed, cellular, sclerotized portion at times interrupted posteriorly. Circuli multiple, there being from 2-5. Legs at times small but of normal composition; claw without denticle or tooth on plantar surface. Antennae 6 - to 7-segmented. Notes. According to Ferris (1950), this genus will probably prove to be of special interest when such genera is Ehrhornia Ferris and Rhodania Goux come under consideration. Tridiscus was named primarily as a part of the process of clarifying the genus Trionymus Berg. To the genus Tridiscus there are now assigned 3 species for North America, 2 of which occur in California.

KEY Species of Tridiscus in North America: Adult Females (Modified from Ferris, 1953)

1.

Only 1 circulus present sporoboli (Cockerell)

FERRIS 2(1).

With at least 2 circuli present 2 Anal ring with anterior two-thirds sclerotized, posterior portion open; with 4 or 5 circuli . . . multiorbis Ferris Anal ring sclerotized throughout; with not more than 3 circuli . distichlii Ferris

Tridiscus distichlii

Ferris

Coastal Grass Mealybug (Figure 189) Trionymus distichlii Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., pp. 69-70, illus. Trionymus distichlii Ferris, 1920. Stanford Univ. Pubs., Univ. Ser., Biol. Sci. 1(1):31. Tridiscus distichlii (Ferris), Ferris, 1950. Atlas of scale insects of North America, Ser. V:249250, illus. Type Locality and Host. Palo Alto (in salt marshes), Santa Clara County, California, on salt grass, Distichlis spicata (Gramineae). North American Distribution. California and Washington. Additional Hosts. Allenrolfea occidentalis (Chenopodiaceae), Cynodon dactylon (Gramineae), and undetermined grasses (Gramineae). External Features and Habitat. According to the original description of this species, the adult female, at maturity, is entirely covered with a fluffy mass of wax and is of a purplish color when divested of secretion. The mealybug is found upon the upper surface of the leaves or in the twig axils. Recognition Characters. Adult females, mounted,

TRIDISCUS Tridiscus Ferris, 1950. Atlas of the scale insects of North America, Ser. V:248. Tridiscus Ferris, 1953. Ibid., Vol. VI:477. Genotype. Trionymus distichlii Ferris. Recognition Characters. Pseudococcidae with ostioles either completely absent or reduced, anterior pair apparently absent, posterior pair very small and detectible only in favorable specimens. Dorsum with cerarii present only on anal lobes, here represented by slender, paired setae. All pores and ducts—trilocular, multilocular, and oral-collar tubulars—of normal form. Anal ring well developed, cellular, sclerotized portion at times interrupted posteriorly. Circuli multiple, there being from 2-5. Legs at times small but of normal composition; claw without denticle or tooth on plantar surface. Antennae 6 - to 7-segmented. Notes. According to Ferris (1950), this genus will probably prove to be of special interest when such genera is Ehrhornia Ferris and Rhodania Goux come under consideration. Tridiscus was named primarily as a part of the process of clarifying the genus Trionymus Berg. To the genus Tridiscus there are now assigned 3 species for North America, 2 of which occur in California.

KEY Species of Tridiscus in North America: Adult Females (Modified from Ferris, 1953)

1.

Only 1 circulus present sporoboli (Cockerell)

FERRIS 2(1).

With at least 2 circuli present 2 Anal ring with anterior two-thirds sclerotized, posterior portion open; with 4 or 5 circuli . . . multiorbis Ferris Anal ring sclerotized throughout; with not more than 3 circuli . distichlii Ferris

Tridiscus distichlii

Ferris

Coastal Grass Mealybug (Figure 189) Trionymus distichlii Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., pp. 69-70, illus. Trionymus distichlii Ferris, 1920. Stanford Univ. Pubs., Univ. Ser., Biol. Sci. 1(1):31. Tridiscus distichlii (Ferris), Ferris, 1950. Atlas of scale insects of North America, Ser. V:249250, illus. Type Locality and Host. Palo Alto (in salt marshes), Santa Clara County, California, on salt grass, Distichlis spicata (Gramineae). North American Distribution. California and Washington. Additional Hosts. Allenrolfea occidentalis (Chenopodiaceae), Cynodon dactylon (Gramineae), and undetermined grasses (Gramineae). External Features and Habitat. According to the original description of this species, the adult female, at maturity, is entirely covered with a fluffy mass of wax and is of a purplish color when divested of secretion. The mealybug is found upon the upper surface of the leaves or in the twig axils. Recognition Characters. Adult females, mounted,

456

TAXONOMY

OF

SPECIES

Fig. 189. Tridiscus distichlii Ferris, collected on salt grass, Distichlis spicata (Gramineae), at Palo Alto (in salt marshes), Santa Clara County, California.

1.95 to 2.50 mm long, 1.00 to 1.40 mm wide; body form elongate oval. Dorsum with anal lobes obsolete. Cerarii present only on anal lobes. Anal lobes cerarius with 2 or 3 elongate, rather stout setae, 2-4 elongate, slender auxiliary setae, very few trilocular pores. Dorsal ostioles represented only by very obscure posterior pair. Trilocular pores very sparsely beset on dorsum. Multilocular disk pores rather numerous, distributed over entire dorsal surface. Oral-collar tubular ducts small, sparsely scattered over dorsum. Body setae short and slender. Anal ring apical with no unusual features; each of its 6 setae but little longer than greatest diameter. Venter with multilocular disk pores fairly numerous in midregion from posterior apical segment of abdomen forward to sixth segment, in lateral areas of abdomen forward to head, scattered on thorax. Trilocular pores relatively few, scattered over venter, certain "clear" areas evident on thorax and head. Oral-collar tubular ducts small, of same shape and size as those on dorsum, scattered over entire venter. Body setae short and slender, sparsely distributed. Normally 3, rarely 4, circuli present, these situated between third and fourth, fourth and fifth, and fifth and sixth segments, all small and circular, the posteriormost usually smaller than others or, at times, absent. The number of circuli seems quite variable ranging from 1 to 3 in specimens examined. Legs very small, though not noticeably stout; hind coxae with cluster translucent pores at base; claw rather slender, without denticle on plantar surface. Antennae 7-segmented, short, moderately stout. Notes. This species is related to Tridiscus multiorbis Ferris, but differs in having the anal ring sclerotized throughout, and usually not more than 3 circuli, whereas in the species last mentioned, only the anterior two-thirds of anal ring is sclerotized, the posterior third open, and there are normally 4 or 5 circuli. California Records (Map 177) Alameda Co.: Oakland, VII-7-32, on Distichlis spicata (B. B. Whitney, CDA). Los Angeles Co.: Lancaster, VII-28-57, on Distichlis spicata (K. C. Stephens, CDA). Orange Co.: Costa Mesa, IX-29-44, on Allenrolfea occidentalis (R. J. Bumgardner, CDA). San Diego Co.: La Jolla, VIII-24-15, on Distichlis sp. (T. D. A. Cockerell, UCD). San Joaquin Co.: Lathrop, IX-17-51, on Distichlis sp. (G. F. Ferris, UCD). San Luis Obispo Co.: Cambria, VI-8-61, on Distichlis spicata (G. G. Beevor, CDA). Santa Barbara Co.: Carpintería, XI-7-32, on Cynodon dactylon (J. F. Weging, CDA). Santa Clara Co.: Palo Alto (in salt marshes), I X 1-16, on Distichlis spicata (G. F. Ferris, UCD). Santa Cruz Co.: Santa Cruz, IX-17-65, on Distichlis spicata (R. F. Wilkey, CDA). Solano Co.: Near Grizzley Island, X-20-59, on Distichlis spicata (R. O. Schuster, UCD). Ventura Co.: Somis, VII-28-61, on Distichlis spicata (W. M. Jones, CDA).

Trinity i

Map 177. Distribution in California of Tridiscus distichlii Ferris.

ElOmdaj

Imp

U S X

Stn BernsrJiño K/vtrùJ* Imprid

Tridiscus multiorbis Ferris Multiple Circulus Mealybug (Figure 190) Tridiscus multiorbis Ferris, 1953. Atlas of scale insects of North America, Vol. VI:478-479, illus. Type Locality and Host. Mouth of Puerto Canyon, near Modesto, Tuolumne County, California, on little bluegrass, Poa sandbergii (Gramineae). North American Distribution. Known only from California. Additional Hosts. Artemisia absinthium (Compositae)—closely associated with Agrostis sp. (Gramineae), Bromus sp. (Gramineae), Hordeum leporinum, and Hordeum spp. (Gramineae). External Features and Habitat. The adult female of this species, has a very small amount of a smooth, white, waxy secretion. The body color is yellow; body shape broadly oval; and legs white. There are no filaments. According to the collectors, no ovisac is produced. This mealybug occurs on the roots of its host. No ants were found in association with the mealybugs. According to Ferris (1953), "All specimens were under leaf sheaths, except those from Hordeum, which occurred about the roots." Recognition Characters. Adult females, mounted, 2.15 to 3.10 mm long, 1.25 to 1.65 mm wide; body form somewhat variable, in fully expanded specimens broadly oval. Dorsal ostioles absent. Dorsum with cerarii represented only by anal lobe pair, these comprising but 2 slender, elongate setae set close together, not acommpanied by auxiliary setae or trilocular pores. Trilocular pores present on dorsum. Multilocular disk pores small, rather numerous on posterior half of body, sparsely distributed on thorax

458

TAXONOMY

OF

SPECIES

Fig. 190. Tridiscus multiorbis Ferris, collected on little bluegrass, Poa sandbergii (Gramineae), at mouth of Puerto Canyon, near Modesto, Tuolumne County, California.

TAXONOMY and head. Dorsum with an occasional oral-collar tubular duct, these ducts being about 3 times as long as their own diameter. Body setae very short and sparse. Anal ring quite distinctive, formed of a sclerotized arc which forms one-half to two-thirds of a circle, this sclerotized area forming anterior and lateral portions of ring, beset with a number of small pores, remainder of anal ring open; each of its 6 setae very small, attaining scarcely more than half greatest diameter of ring. Venter with multilocular disk pores distributed similar to those on dorsum, there being only a slight concentration about vulva, wide submarginal band along body, crowded group about spiracles. Trilocular pores present. Oral-collar tubular ducts very few, distributed similar to those on dorsum. Body setae short and slender, very sparsely scattered. Circuli very small, variable in size, when seen in profile they seem to form a deep little cup with strongly sclerotized walls, situated on each of apparent 4 segments of abdomen (occasional specimen may have a fifth circulus). Legs very small, but completely developed; hind femur and tibia with few translucent pores; claw without denticle on plantar surface. Antennae usually 6-segmented, occasionally 7-segmented. Notes. Ferris states: "Chiefly because of the difference in body form this species does not at first glance look much like the type of its genus, yet there appears to be nothing except the presence of the additional circulus and the peculiar form of the anal ring by which it could be separated into another genus. It seems actually to belong with Tridiscus." Some of the specimens taken on grass at Asilomar, Pacific Grove, California, have 5 circuli instead of 4, although they agree in detail with the others and are here considered to represent T. multiorbis. Tridiscus multiorbis resembles T. distichlii (Ferris), but differs from it in having the anal ring with only the anterior two-thirds sclerotized, the posterior third open, and in having 4 or 5 circuli, whereas in the species last mentioned the anal ring is sclerotized throughout, and there are not more than 3 circuli. California Records (Map 178) Kern Co.: Near Simmler, III—29—35, on Poasandbergii (G. F. Ferris, UCD). Los Angeles Co.: Near Lancaster, IV-27-36, on "undetermined grass" (G. F. Ferris, UCD); I V 6-64, on "grass" (roots) (G. G. Beevor and J. L.

SPECIES

459

Johnson, CDA); Lancaster (Neenach), IV-4-61, on Hordeum leporinum (roots) (G. G. Beevor, CDA); Van Nuys (San Fernando Valley), IH-21-33, on "grass" (roots) (L. E. Myers, LEM). Monterey Co.: Sand dunes near Asilomar, IV5-20, on "undetermined grass" (G. F. Ferris, UCD); Parkfield, 111-29-35, on Poa sandbergii (G. F. Ferris, UCD). Riverside Co.: 3.4 miles northeast Gilman Springs, III-9-63, on Artemisia absinthium (D. R. Miller, UCD); 1 mile north White Water, 1-25-65, on Bromus sp. (crown) (D. R. Miller, UCD). San Bernardino Co.: Morongo Valley, IV-15-65, on Bromus sp. (lower crown) (D. R. and J. F. Miller, UCD). San Luis Obispo Co.: Carrizo Plain, 111-30-35, on Hordeum Sp. (G. F. Ferris, UCD). Tuolumne Co.: Puerto Canyon, near Modesto, III—15—35, on Poa sandbergii (G. F. Ferris, UCD).

TRIONYMUS Westwoodia Signoret, 1875. Ann. Soc. Ent. de France (Ser. 5) 4:337 (preoccupied). Signoretia Kraatz, 1888. Deut. Ent. Ztschr. 32:176 (preoccupied). Bergrothia Kraatz, 1888. Ibid. 32:360 (preoccupied). Bergrothiella Reitter, 1898. Wien. Ent. Ztg. 17: 54-56 (preoccupied).

OF

BERG

Pergandiella Cockerell, 1899. Proc. Acad. Nat. Sci. Phila., p. 266. Trionymus Berg, 1899. Buenos Aires Mus. Nac. Comunicaciones 1:77-80. Trionymus Berg, Fernald, 1903. Mass. Agr. Expt. Sta. Spec. Bull. 88, p. 96. Pergrolhula Strand, 1926. Arch. f. Naturgesch. 92 (Abt. A, h. 8) :47.

460

TAXONOMY

OF

SPECIES

Trionymus Berg, Zimmerman, 1948. Insects of Hawaii, Univ. Hawaii Press, 5:256-257. Trionymus Berg, Ferris, 1950. Atlas of scale insects of North America, Ser. V:251. Trionymus Berg, Ferris, 1953. Ibid., Vol. VI:482. Trionymus Berg, McKenzie, 1960. Hilgardia 29 (15):764. Trionymus Berg, Williams, 1962. Bull. Brit. Mus. (Nat. Hist.) Ent. 12(1) :58. Trionymus Berg, De Lotto, 1964. Ibid. 14(8):386. There is some question as to whether Trionymus or Pergandiella has priority, but Trionymus was adopted by the Fernald Catalogue, and several authors, including Ferris (1950-1953), McKenzie (1960), Williams (1962), and De Lotto (1964), have recognized it as valid. Genotype. Westwoodia perrisii Signoret. Recognition Characters. Pseudococcidae, for the most part, with body form at maturity elongate oval. Two pairs of dorsal ostioles. Cerarii reduced in number; occasionally a frontal pair present, those along side of body absent, leaving from 1-4 pairs at end of abdomen, these always including anal lobe pair, total of 5 or fewer pairs of cerarii, including frontal ones. Oral-rim tubular ducts absent anywhere on body. Multilocular disk pores present or absent on dorsum. Anal ring of normal cellular development and position, never situated posterior to anal lobe cerarii, bearing 6 setae each approximately twice as long as diameter of ring. Venter always with at least a few multilocular disk pores. Circulus present or absent, when present usually small, circular or oval, at times quadrate-shaped, at times divided by intersegmental line, though usually not. Legs normally developed, tarsal claw without denticle on plantar surface. Antennae 6 - to 8-segmented. The majority of the species live on grasses. Notes. This is one of the very oldest genera of the Pseudococcidae. This genus is rather closely related to Dysmicoccus Ferris. Ferris (1950) placed considerable emphasis on the shape of the circulus which, in Trionymus, is usually small, round or oval, whereas in Dymicoccus it is normally quite large and divided by intersegmental fold. I (I960) disregarded the shape of the circulus since certain species were subsequently found in which this structure was somewhat quadrate in appearance and obviously divided by the intersegmental line. Trionymus, as here interpreted, has but 5 or fewer pairs of cerarii, including frontal ones, lacks oral-rim tubular ducts, and the majority of the species involved are to be found on grasses. Four new California species of Trionymus are described in this book. To the genus there are now assigned 20 species for North America, 15 of which occur in California.

KEY Species of Trionymus in North America: Adult Females

1. 2(1). 3(2).

4(3).

5(2).

6(1). 7(6).

8(7).

9(8).

Circulus present 6 Circulus absent 2 Cerarii present only on anal lobes . . . . 3 Cerarii present on last 2 abdominal segments 5 Ventral multilocular disk pores present along submargin below eye and near spiracles winnemucae McKenzie Ventral multilocular disk pores absent along submargin below eye and near spiracles 4 Discoidal pores with poriferous centers numerous on both body surfaces; hind coxae not perceptibly larger than coxae of other legs nanus Cockerell Discoidal pores absent anywhere on body; hind coxae definitely larger than coxae of other legs smithii (Essig) Hind tibiae slender, with at most only faint trace of inconspicuous translucent pores near apical half of segment; antennae 8-segmented; dorsal oral-collar tubular ducts small, all approximately same size furvus McKenzie Hind tibiae robust, swollen, with numerous conspicuous translucent pores situated over most of segment; antennae 9-segmented (rarely 8-segmented); a few comparatively large dorsal oral-collar tubular ducts, associated with numerous smaller ducts of similar type myersi McKenzie Cerarii present only on anal lobes . . . 7 With more than 1 pair of cerarii present 10 Antennae normally 7-segmented; dorsal multilocular disk pores present on mesothorax forward to head 8 Antennae normally 8-segmented, rarely 7-segmented; dorsal multilocular disk pores absent on mesothorax forward to head, except for an occasional pore in this area . utahensis (Cockerell) Ventral multilocular disk pores on abdomen unusually numerous, those of segments 5 to 7, medially, arranged in broad bands of 4 pores deep . . . . clandestinus McConnell Ventral multilocular disk pores on abdomen, medially, in bands nowhere more than 2 pores deep 9 Body form of adult female narrowly oval caricis McConnell Body form of adult female broadly pyri-

TAXONOMY

10(6).

11(10).

12(11).

13(12).

14(10).

15(14).

16(15).

17(15).

18(17).

19(14).

form . . . . magnus (Cockerell and Cockerell) Circulus normally quadrate-shaped, often divided by segmental line of segments between which it lies 11 Circulus small, circular or oval, not divided by segmental line of segments between which it lies 14 Dorsal multilocular disk pores present at least on last 3 or 4 abdominal segments 12 Dorsal multilocular disk pores absent on abdomen modocensis (Ferris) Ventral multilocular disk pores present on thorax 13 Ventral multilocular disk pores absent on thorax haancheni McKenzie Frontal cerarius present frontalis McKenzie Frontal cerarius absent quadricirculus McKenzie Cerarii definitely recognizable only on last 2 abdominal segments 15 Cerarii definitely recognizable on more than last 2 abdominal segments . . . 19 Anal lobe cerarius with a distinct surrounding sclerotized area 16 Anal lobe cerarius without a surrounding sclerotized area 17 Multilocular disk pores present over entire dorsum claviger (King and Tinsley) Multilocular disk pores, if present on dorsum at all, confined to abdominal segments americanus (Cockerell) Ventral multilocular disk pores very numerous in lateral areas of both thorax and abdomen 18 Ventral multilocular disk pores absent or very few in lateral areas of both thorax and abdomen violascens Cockerell Dorsal multilocular disk pores numerous on all segments of abdomen; antennae 8-segmented; occurs on Agavaceae, mainly Phormium diminutus (Leonardi) Dorsal multilocular disk pores restricted to last 2 abdominal segments, except for occasional pore along submargin of segments 6 and 7; antennae 7-segmented; occurs on Gramineae festucae (Kuwana) Antennae 7-segmented; ventral multilocular disk pores present on thorax, especially in lateral areas; anal lobe cerarius without surrounding sclerotized area mocus Ferris Antennae 8-segmented; ventral multilocular disk pores absent on thorax, rarely an occasional pore; anal lobe

OF SPECIES

461

cerarius surrounded by distinct sclerotized area dolus Ferris

Trionymus americanus (Cockerell) American Grass Mealybug (Figure 191) Pergandiella americana Cockerell, 1899. Proc. Acad. Nat. Sci. Phila., pp. 266-268. Trionymus americanus (Cockerell), Fernald, 1903. Mass. Agr. Expt. Sta. Spec. Bull. 88, p. 96. Trionymus americanus (Cockerell), Ferris, 1950. Atlas of scale insects of North America, Ser. V:253-254, illus. Type Locality and Host. Washington, D. C. (on grounds of Department of Agriculture), on ash, Fraxinus sp. (Oleaceae). North American Distribution. California, Maryland, Mississippi, New York, Utah, and Washington, D. C. Additional Hosts. Cydonia sp. (Rosaceae), Elymus glaucus, E. virginicus (Gramineae), and undetermined grasses (Gramineae). External Features and Habitat. According to Cockerell (1899), this mealybug is very slender, of a brownish-red color, and covered with a mealy excretion giving it a grayish or moldy appearance. Prior to egg laying, a woolly excretion is formed which is rather sparse anteriorly, but becomes very dense toward posterior end of body. A pale purplishcolored ovisac is formed. This mealybug was reported as occurring in the cracks on the trunk of the host. The species belongs to a grass-infesting group, and it is believed that the records from ash and later from beneath paper bands on quince were occasioned by the habit of crawling around before forming the ovisac. It is assumed that the species occurs beneath the leaf sheaths of its host. Recognition Characters. Adult females, mounted, 3.10 to 3.60 mm long, 1.20 to 1.60 mm wide; body form unusually elongate oval. Dorsum with cerarii present only on last 2 abdominal segments. Anal lobe cerarius with 2 moderately stout, conical setae, a number of slender auxiliary setae of various lengths, accompanied by scattered trilocular pores, all contained within oval area normally quite well sclerotized. Penultimate cerarius with 2 slightly smaller conical setae, 3-4 slender auxiliary setae, small cluster trilocular pores, no basal sclerotization. Trilocular pores numerous, evenly beset on dorsum. Minute circular pores scattered. Multilocular disk pores usually present from a few to fairly numerous on last 3 or 4 abdominal segments. Oral-collar tubular ducts with a slight oral collar present in transverse series on abdominal segments, sparsely scattered over thoracic and head regions. Body setae slender, sparsely beset on dorsum. Anal ring apical

462

TAXONOMY

OF

SPECIES

Fig. 191. Trionymus americanus (Cockerell), collected on "lawn grass" (Gramineae), at Alturas, Modoc County, California. (Illustration prepared from specimens collected on quince, at Ringgold, Maryland.)

T AXON

with no unusual features; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores present in midregion of abdomen from posterior apical segment forward to fifth segment. Trilocular pores evenly beset on venter. Minute circular pores scattered. Oralcollar tubular ducts, similar to those on dorsum, in considerable numbers in lateral areas from eighth abdominal segment forward to head, sparsely distributed in midregions, especially so on abdomen. Body setae slender, sparsely distributed on venter. Circulus slightly variable in size, normally quite small and typical for members of this genus. Legs well developed; hind coxa with small cluster translucent pores at base; claw without denticle on plantar surface. Antennae normally 8-segmented. Notes. This species is rather closely related to Trionymus haancheni McKenzie, differing from it mainly in the circulus which is quite small, circular or oval, not folding along intersegmental line of the segments between which it lies; and in the presence of ventral multilocular disk pores on the fifth abdominal segment. Trionymus haancheni, on the other hand, has a comparatively large circulus, normally quadrateshaped, usually divided by segmental line; and the ventral multiloculars are absent on the fifth abdominal segment. California Records (Map 179) Alpine Co.: Markleeville, Pleasant Valley, X-5-65, on grass (roots) (R. F. Wilkey, CDA). Modoc Co.: Alturas, IX-5-51, on "lawn grass" (L. White, CDA). San Diego Co.: Rancho Sante Fe, VII-10-58, on "grass" (S. M. Klopfer, CDA). Santa Cruz Co.: Santa Cruz, X-20-40, on "lawn grass" (H. Washburn, CAS).

Map 179. Distribution in California of Trionymus

(Cockerell).

americanus

OMY

OF

SPECIES

463

Trionymus caricis McConnell Carex Mealybug (Figure 192) Trionymus caricis McConnell, 1941. Proc. Wash. Ent. Soc. 43(5):97-99, illus. Trionymus caricis McConnell, Ferris, 1950. Atlas of scale insects of North America, Ser. V: 257-258, illus. Trionymus caricis McConnell, McKenzie, 1964. Hilgardia 35(10): 270. Type Locality and Host. Bowie, Prince Georges County, Maryland, on sedge, Carex tribuloides (Cyperaceae). North American Distribution. California, Maryland, and Tennessee. Additional Hosts. Ely mus triticoides (Gramineae), Unióla ornithorhynca (Gramineae); undetermined grasses (Gramineae); and foliage of certain undetermined plants. External Features and Habitat. According to the original description, the body of the adult female of this species is pinkish to reddish in color, lightly dusted over with powdered wax. An ovisac is formed of indefinite character on account of position on host plant, but usually it is long, narrow, and flat. The mealybug occurs beneath the leaf sheaths of the host. Recognition Characters. Adult females, mounted, 2.00 to 3.00 mm long, 0.90 to 1.00 mm wide; body form slender, more or less parallel-sided. Dorsum with cerarii present only on anal lobes, these with 2 quite small, conical setae, accompanied by 3-4 slender auxiliary setae of varying lengths, very few trilocular pores, no basal sclerotization. Trilocular pores rather sparsely distributed on dorsum. Multilocular disk pores variable in number, occurring on last 5-6 abdominal segments, very few situated along body margin on anterior abdominal segments forward to head. Oral-collar tubular ducts variable in number, ranging from a few to fairly numerous in rows on abdominal segments, scattered on thorax and head; orifice diameter of ducts slightly greater than trilocular pore. Body setae small, slender, scattered on dorsum. Anal ring apical, with no unusual features; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores present from posterior apical segment of abdomen forward to head, for most part scattered, occurring in more or less distinct bands on abdominal segments 8-5, and in lateral areas of all abdominal segments. Trilocular pores fairly evenly beset on venter. Oral-collar tubular ducts of 2 sizes occur in small numbers in midregion of most abdominal segments and in lateral areas forward to head. Ventral body setae quite small, slender. Circulus quite small, circular or oval, though situated on segmental line apparently not divided by it. Legs quite small, not especially stout; hind coxae with cluster trilocular pores at base; claw without denticle on plantar surface. Antennae normally 7-segmented, although at times with 6, or even 8, segments.

464

TAXONOMY

OF

SPECIES

Fig. 192. Trionymus caricis McConnell, collected on wild rye, Elymus triticoides (Gramineae), at Patterson, Stanislaus County, California. (Illustration prepared from type material.)

TAXONOMY Notes. This species is very closely related to Trionymus utahensis (Cockerell), differing principally in possessing multilocular disk pores which are scattered over entire dorsum, and the antennae which is normally 7-segmented (infrequently 6- or 8-segmented). Trionymus utahensis, on the other hand, has multilocular disk pores usually confined to last 4 or 5 abdominal segments (entirely absent on mesothorax forward to head), and the antennae are normally 8-segmented. California Records (Map 180) Plumas Co.: 6 miles east Quincy, VII-9-64, on Elymus glaucus (leaf sheaths) (D. R. Miller, UCD). Siskiyou Co.: Near Dunsmuir (Bradley Mountain), IX-20-61, on "undetermined plants" (R. O. Schuster, UCD). Stanislaus Co.: Patterson, summer, 1964, on Elymus triticoides (J. Azevado, CDA).

Trionymus diminutus (Leonardi) New Zealand Flax Mealybug (Figure 193) Pseudococcus diminutus Leonardi, 1918. Portici R. Scuola Super. Agr. Lab. Zool. Gen. e Agr. Boi. 12:198-201, illus. Trionymus calceolariae (Maskell), Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., p. 67-68 (as a misidentification). Pseudococcus diminutus Leonardi, 1920. Portici, Della Torre, pp. 397-399, illus. Trionymus diminutus (Leonardi), Morrison, 1925. Jour. Agr. Res. 31 (5) :495-496, illus.

OF

SPECIES

465

Trionymus diminutus (Leonardi), Ferris, 1950. Atlas of scale insects of North America, Ser. V:261262, illus. Type Locality and Host. Bordighere, Liguria, Italy, on New Zealand flax, Phormium tenax (Agavaceae). North American Distribution. Known only from California. Additional Hosts. According to Morrison (1925), "So far as positively known, the species is normally confined to Phormium as a host, although, as pointed out in discussing the intricacies of Maskell's utilization of the name calceolariae, the specimens recorded by him from Cordyline australis (Agavaceae) in 1893 may reasonably be given a tentative assignment here." Other tentative host determinations associated with certain collections of this mealybug include: A gave sp. (Agavaceae), Cordyline australis (Agavaceae), Fraxinus ? sp. (Oleaceae), and Yucca sp. (Agavaceae). External Features and Habitat. This mealybug is covered with a fine, mealy, wax secretion. A pair of short, curved, caudal filaments are present, and the lateral ones, if present, are very small and inconspicuous. A fluffy waxy ovasic is formed that almost completely hides the insect. This species infests the plant portions above ground. Recognition Characters. Adult females, mounted, 2.60 to 5.50 mm long, 1.40 to 2.90 mm wide; body form rather elongate, broadly oval. Dorsum with cerarii present on last 2 abdominal segments, or at times with suggestion of antepenultimate cerarius on seventh segment. Anal lobe cerarius with 2 quite short, stout, usually slightly acorn-shaped setae, essentially no auxiliary setae, almost no accompanying concentration trilocular pores; penultimate cerarius with either 2 somewhat more slender and more conical setae or, at times, 1 such seta and a longer, slender seta, no auxiliary setae or accompanying trilocular pores. Occasionally cerarius of seventh segment indicated by pair slender setae set close together. Trilocular pores numerous, evenly beset on dorsum. Minute circular pores scattered. Multilocular disk pores abundant over entire dorsum, especially in lateral areas and across abdominal segments. Oralcollar tubular ducts small, situated in considerable numbers over dorsum, especially in lateral areas, these ducts with orifices not greater in diameter than a trilocular pore, and scarcely more than twice as long as wide. Body setae quite numerous, of various lengths, some moderately long and stout. Anal ring apical, normal cellular appearance; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores extremely abundant on posterior apical segment of abdomen, in a broad band along posterior border of eighth and seventh segments, less abundant on sixth segment, also abundant in lateral areas of all segments forward to prothorax, few on head. Trilocular pores numerous, evenly beset on venter. Minute circular pores

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Fig. 193. Trionymus diminutus (Leonardi), collected on New Zealand flax, Phormium tenax (Agavaceae), at Berkeley, Alameda County, California.

TAXONOMY

scattered. Oral-collar tubular ducts of same size and shape as those of dorsum fairly numerous especially in lateral areas. Body setae of various lengths beset on venter, some of those near margins quite short. Circulus small and circular. Legs well developed, not especially stout; hind coxa with cluster of translucent pores at base; claw without denticle on plantar surface. Antennae 8-segmented, slender. Notes. This species was for many years misidentified as Pseudococcus calceolariae (Maskell). The confusion was clarified by Morrison (1925), who recorded that the species associated with Phormium is apparently that described by Leonardi (1918) as Pseudococcus diminutus. According to Ferris (1950), who prepared the accompanying illustration, there is some variation in diminutus, the cerarian setae of the anal lobe pair at times not being as definitely acorn-shaped as is here illustrated, and the penultimate cerarii at times having 1 conical and 1 slender seta, but these variations are not such as to cause any special difficulty in recognizing the species. This species is close to Trionymus festucae (Kuwana), but differs in possessing numerous dorsal multilocular disk pores on all segments of abdomen, whereas in T. festucae these structures are restricted to the last 2 abdominal segments, except for an occasional pore along submargin of segments 6 and 7. California Records (Map 181)

Alameda Co.: Berkeley, X-2-61, on Agave sp. (U. Nur, UCD); Berkeley (University of California Campus), IX-12-06, on Phormium tenax (O. E. Bremner, UCD); VIII-19-14, on Phormium tenax (C. P.

OF

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467

Clausen, UCD); 11-13-17, on Phormium tenax (E. O. Essig, CAS); Berkeley, IX-16-11, on Yucca sp. (G. A. Coleman, UCD); Fremont, V-9-62, on Fraxinus ? sp. (M. Hansen, CDA); Niles, 111-14-57, on Phormium sp. (F. H. Duffie, CDA); V-7-58, on Phormium sp. (F. H. Duffie, CDA); IV-18-56, on Phormium tenax (P. Sweigart and E. Henning, CDA). Marin Co.: Corte Madera (quarantined at San Francisco), 11-24-55, on Phormium sp. (R. Bazzini and D. J. Bingham, CDA); Fairfax, IV-1-52, on Phormium tenax (R. P. Allen, CDA). San Francisco Co.: San Francisco, VII-12-41, on Phormium tenax (M. R. Bell, CDA). San Mateo Co.: Redwood City (nursery), 1-10-50, on Phormium tenax (R. P. Allen, CDA); Portolo Valley, VIII-18-65, on Phormium tenax (F. E. Sampson, CDA). Santa Clara Co.: San Jose, IV-20-64, on Phormium tenax (E. C. Winkler and C. O. Howe, CDA); III—11-60, on Phormium sp. (E. C. Winkler and G. F. Prole, CDA).

Trionymus dolus Ferris Dolus Mealybug (Figure 194; Color Plate XXV) Trionymus dolus Ferris, 1950. Atlas of scale insects of North America, Ser. V: 263-264, illus. Type Locality and Host. Ravenswood, now called north Palo Alto, Santa Clara County, California, on wild rye, Elymus sp. (Gramineae). North American Distribution. California, Oregon, and Washington. Additional Hosts. Cynodon dactylon (Gramineae), Elymus condensatus (Gramineae), and Elymus glaucus? (Gramineae). External Features and Habitat. The adult female of this species is light yellow-cream colored and possesses wax filaments developed on last 3 abdominal segments, those of anal lobes being quite stout and nearly one-third the length of the body. A loose amorphous ovisac is produced at maturity, although this covers but little of the insect. The eggs are light yellow when first laid (see col. pi. XXV). The species is found beneath the sheathing bases of the leaves or, at times, exposed upon the leaf surface. Recognition Characters. Adult females, mounted, 1.95 to 5.00 mm long, 1.00 to 2.50 mm wide; body form elongate oval. Dorsum with cerarii normally present on last 3 abdominal segments, also a frontal pair. Anal lobe cerarius with 2 moderately large, conical setae, as many as 10 auxiliary setae of various lengths, quite numerous but scattered trilocular pores, all borne upon quite well-defined, oval area that tends to be definitely sclerotized; penultimate cerarius with 2 somewhat smaller, conical setae, 5-6 slender auxiliary setae, cluster trilocular pores, no sclerotized base; antepenultimate cerarius with 2 slender conical setae, very slight concentration trilocular pores, or

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Fig. 194. Trionymus dolus Ferris, collected on wild rye, Elymus sp. (Gramineae), at Ravenswood, now called north Palo Alto, Santa Clara County, California.

COLOR PLATE XXV. Dolus mealybug, Trionymus dolus Ferris, in leaf-sheaths of giant ryegrass, Elymus condensatus (Gramineae), at Bel Air, Los Angeles County, California.

these setae slender and elongate, no auxiliary setae or basal sclerotization; cerarius of sixth abdominal segment composed merely of a pair of slender setae, or these at times obscure, not developed. Frontal cerarius composed merely of a pair of small, conical setae, small group trilocular pores, no sclerotization. Trilocular pores evenly beset on dorsum. Minute circular pores scattered. Multilocular disk pores either absent or in small numbers scattered in submedian area on last 3 or 4 abdominal segments, an occasional pore near lateral margins on these and other segments, infrequently on head. Oral-collar tubular ducts small, orifices slightly less in diameter than a trilocular pore, perhaps 4 times as long as wide, quite abundant over dorsum of abdomen and in lateral areas of thorax and head. Body setae small and slender, variable in size. Anal ring normal for genus; each of its 6 setae about twice as long as diameter of ring. Venter with multilocular disk pores fairly numerous, occurring in an irregularly single or double row along posterior borders of abdominal segments 8-5, in area posterior to vulva, occasional pore in lateral areas, absent elsewhere on venter. Trilocular pores evenly beset on venter. Minute circular pores scattered. Oral-collar tubular ducts of same size and shape as those on dorsum situated in small numbers in midregions of abdominal segments, more abundant in lateral areas of these segments, sparsely distributed in lateral areas of thorax and head. Body setae slender, variable in length, a few longer than those on dorsum. Circulus moderately large, circular. Legs well developed; hind coxa with few translucent pores at base; hind tibia with similar pores scattered along segment; claw without denticle on plantar surface. Antennae normally 8-segmented, slender. Notes. This species is similar to Trionymus mocus Ferris, but may be separated from it by 8-segmented antennae, absence of ventral multilocular disk pores in lateral areas of thorax and on head, and anal lobe cerarius surrounded by distinct area of sclerotization. Trionymus mocus, on the other hand, possesses 7-segmented antennae, has ventral multilocular disk pores in lateral areas of thorax and on head, and anal lobe cerarius is without surrounding sclerotization. California Records (Map 182) Los Angeles Co. : Bel Air, VI-20-64, on Elymus condensatus (M. Kosztarab and A. Thiery, UCD); VIII-7-64, on Elymus condensatus (M. F. Benson, UCD). Monterey Co.: Pacific Grove, XII-1-17, on Elymus glaucus ? (G. F. Ferris, UCD). San Diego Co.: Rancho Santa Fe, VII-7-61, on Cynodon dactylon (E. I. Schlinger, UCD). Santa Barbara Co.: Montecito, 1-26-65, on Elymus sp. (G. G. Beevor and E. F. Nickerson, CDA). Santa Clara Co.: Ravenswood (now North Palo Alto), X-19-?, on Elymus sp. (collector ?, UCD). Ventura Co.: Ojai (Dennison Grade), VI-10-58, on Elymus sp. (L. R. Gillogly, Weber, and Holmer, CDA).

Trionymus festucae (Kuwana) North Coast Mealybug (Figure 195) Ripersia festucae Kuwana, 1901. Proc. Cal. Acad. Sci. (Ser. 3, Zool.) 2(10) : 401-402, illus. Ripersia festucae Kuwana, Cockerell, 1902. Ann. Mag. Nat. Hist. (Ser. 7) 10:21. Trionymus festucae (Kuwana), Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., p. 70. Trionymus festucae (Kuwana), Ferris, 1950. Adas of scale insects of North America, Ser. V:265266, illus. Type Locality and Host. Black Mountain near Stanford University, Santa Clara County, California, on fescue, Festuca scabrella (Gramineae). North American Distribution. California and Louisiana. Additional Hosts. Anthoxanthum odoratum (Gramineae), Deschampsia ceaspitosa (Gramineae), and unidentified grass (Gramineae). The record from "wild strawberry" is in all probability due to the habit of wandering about before oviposition. External Features and Habitat. The adult female of this species is elongate-elliptical in form, and is lightly covered with a white, waxy secretion. The body is pink to purple, and the legs cream-white in color. No filaments were observed. An elongate filamentous ovisac is formed at maturity which is approximately twice as long as body. The ovisac covers only a small portion of the abdomen, most of it being posterior to body. Some parasitism was noted on this mealybug, apparently by a Dipterous larva. The mealybug occurs concealed beneath the sheathing bases of grass leaves. Recognition Characters. Adult females, mounted, 1.95 to 3.50 mm long, 0.80 to 1.20 mm wide; body form elongate oval.

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Fig. 195. Trionymus festucae (Kuwana), collected on fescue, Festuca scabrella (Gramineae), at Black Mountain near Stanford University, Santa Clara County, California.

Snkiifn Dorsum with cerarii present on last 2 abdominal segments. Anal lobe cerarius with 2 rather slender, conical or slightly lanceolate setae, 2-3 slender auxiliary setae, no noticeable concentration trilocular pores, slight sclerotization about their bases. Penultimate cerarius with 2 slightly smaller setae of same type, unaccompanied by auxiliary setae or concentration of trilocular pores. Trilocular pores rather evenly beset on dorsum. Multilocular disk pores in small numbers across abdominal segments 8-9, occasional pore of this type near lateral margins of other segments, or even on head. Oral-collar tubular ducts very short and broad, their diameter somewhat more than that of a trilocular pore, their length only slightly greater than diameter, scattered across all abdominal segments, less numerous on thorax and head. Body setae sparse, small, and slender. Anal ring with no unusual features; each of its 6 setae nearly twice as long as diameter of ring itself. Venter with multilocular disk pores in large numbers from posterior apical segment of abdomen forward to sixth segment in both midregion and lateral areas, in loose clusters in lateral areas of all other segments forward to head. Trilocular pores sparsely beset on venter. Oral-collar tubular ducts of same type as those on dorsum mingled with lateral clusters of multilocular disk pores, also situated across midregion of all abdominal segments from apical forward at least to fifth. Body setae of various lengths, some slightly longer than those on dorsum. Circulus very small, circular. Legs well developed, rather small; hind coxae with cluster translucent pores at base; claw without denticle on plantar surface. Antennae normally 7-segmented. Notes. This species is related to Trionymus diminutus (Leonardi), but differs in having very few dorsal multilocular disk pores restricted mostly to last 2 abdominal segments, normally a 7-segmented antenna, and occurring only on Gramineae, whereas, in the species last mentioned, the dorsal multilocular disk pores are extremely numerous on all segments of abdomen, antenna is normally 8-segmented, and it is known only from Agavaceae, mainly on Phormium. California Records (Map 183) Humboldt Co.: Big Lagoon, VIII-12-63, on Deschampsia ceaspitosa (D. R. Miller, UCD). Marin Co.: 7 miles east Nicasio, VIII-26-63, on "grass" (D. R. Miller, UCD). Mendocino Co. : 4 miles north Point Arena, VIII9-63, on Anthoxanthum odoratum (D. R. Miller, UCD). San Mateo Co.: Near Woodside, IV-28-17, on "wild strawberry" (G. F. Ferris, UCD). Santa Clara Co.: Black Mountain near Stanford University, 111-24-1900, on Festuca scabrella (S. I. Kuwana, UCD). Sonoma Co.: Fort Ross, IX-1937, on ? (G. F. Ferris, UCD); 8 miles south Stewarts Point, VIII8-63, on Deschampsia caespitosa (D. R. Miller and J. S. Buckett, UCD).

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Trionymus frontalis McKenzie, new species Frontal Grass Mealybug (Figure 196) Type Locality and Host. Chatsworth, Los Angeles County, California, on wild rye, Elymus sp. (Gramineae), collected July 9, 1958, by L. R. Gillogly. This represents the only known collection of this species. Type Material. Holotype adult female (single specimen on slide) in collection, California Department of Agriculture, Sacramento. Paratype adult females deposited in museum, University of California, Davis. North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. No available information regarding external appearance of this mealybug. It is assumed that the species infests the leaf-blade sheaths of its grass host. Recognition Characters. Adult females, mounted, 3.00 to 4.80 mm long, 1.50 to 2.20 mm wide; body form elongate oval. Dorsum with cerarii present on last 2 abdominal segments and a frontal pair on head. Anal lobe cerarius with 2 rather small, conical setae, several slender auxiliary setae, fair concentration trilocular pores, no basal sclerotization. Penultimate cerarius with 2 slightly smaller conical setae, fewer slender auxiliary setae, slight concentration trilocular pores, no basal sclerotization. Frontal cerarius on head similarly constructed except for perhaps fewer slender auxiliary setae and associated trilocular pores. Trilocular pores extremely numerous and evenly beset on dorsum. Multilocular disk pores rather

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Fig. 196. Trionymus frontalis McKenzie, new species, collected on wild rye, Elymus sp. (Gramineae), at Chatsworth, Los Angeles County, California.

sparse, scattered on last 4 or 5 abdominal segments, 1 or 2 on metathorax, absent elsewhere on dorsum. Minute circular pores or modified discoidal pores, variously shaped, scattered in numbers over entire dorsal surface. Oral-collar tubular ducts of 2 sizes present; larger-sized ones, with very short tubes, extremely numerous over whole surface, perhaps as abundant as trilocular pores; smaller-sized pores scattered over most of dorsum. Body setae slender, variable in length, rather abundant over dorsum. Anal ring apical, normal in form and size for members of this genus; each of its 6 setae about twice as long as diameter of ring. Venter with multilocular disk pores especially numerous in midregion of last 3 or 4 abdominal segments, scattered and less abundant on segment 5 forward to thorax, few distributed in sternal region of thorax, several noticed on head. Trilocular pores abundant, fairly evenly beset on venter except for certain "clear" areas on thorax. Minute circular pores, or modified discoidal pores, similar to those on dorsum, distributed over entire ventral surface. Oral-collar tubular ducts of 2 sizes, resembling those on dorsum and somewhat similarly distributed. Body setae slender, quite numerous, of various lengths, for most part slightly longer than those on dorsum. Circulus present, transversely oval, divided by segmental line across which it lies. Legs moderately large; hind coxae with translucent pores at base; very few such pores distributed along tibial segment; claw without denticle on plantar surface, or with but very slight swelling in this area. Antennae 8-segmented. Mouthparts moderately broad. Notes. This species seems somewhat related to Trionymus quadricirculus McKenzie, herein described as new, but differs in possessing definite frontal cerarii on head, and dorsal multilocular disk pores absent on fourth abdominal segment, whereas, in the species last mentioned, there are no frontal cerarii on head, and at least a few dorsal multilocular disk pores present on fourth abdominal segment. In addition, the longest dorsal body setae of T. frontalis are almost twice as long as those of T. quadricirculus. California Records (Map 184) Los Angeles Co.: Chatsworth, VII-9-58, on Elymus sp. (L. R. Gillogly, C D A ) .

Trionymus furvus McKenzie, new species Dusky Grass Mealybug (Figure 197) Type Locality and Host. 15 miles north of Bishop, Inyo County, California, on wild rye, Elymus triticoides (sheaths), July 17, 1963, D. R. Miller collector. Type Material. Holotype adult female (single specimen on slide) and female paratypes in museum, University of California, Davis. Additional adult female paratypes in United States National Museum, Washington, D. C., and in collection of California State Department of Agriculture, Sacramento.

North American Distribution. Known only from California. Additional Hosts. None. External Features and Habitat. This mealybug is elongate oval, lightly covered with a powdery, gray, wax secretion, and with a pair of moderately long caudal filaments. The body color is dark green to nearly black (hence the specific name, furvus.) A posterior ovisac is formed covering nearly all of the abdomen. Construction of this sac appears to begin before the insects become fully adult. The mealybug is apparently ovoviparious. The species occurs beneath the leaf sheaths of its grass host and on the grass blades. A very severe infestation was noted with never fewer than 100 specimens on each plant. The root area was found entirely free of this mealybug. Recognition Characters. Adult females, mounted, 1.98 to 2.50 mm long, 1.00 to 1.15 mm wide; body form elongate, more or less parallel-sided. Dorsum with cerarii present on last 2 abdominal segments. Anal lobe cerarius with 2 rather slender, conical setae, 2 - 3 slender auxiliary setae, slight concentration trilocular pores, no basal sclerotization. Penultimate cerarius variable, containing either 2 somewhat more slender, conical setae, one such seta and a longer, slender seta, or both setae long and slender, all with 0 - 1 smaller, slender auxiliary seta, scarcely any associated trilocular pores, no basal sclerotization. Trilocular pores evenly beset on dorsum. Multilocular disk pores occurring in small numbers on abdominal segments, less numerous in midregion, very sparsely scattered on thorax. Oral-collar tubular ducts short, about twice as long as wide, with orifices as

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Fig. 197. Trionymus furvus McKenzie, new species, collected on wild rye, triticoides (Gramineae), north of Bishop, Inyo County, California.

Elymus

great in diameter as trilocular pores, in irregular rows across abdominal segments, fairly numerous, though scattered, on thorax and head. Body setae slender, of various lengths, some noticeably longer than those on dorsum. Anal ring apical or but slightly removed from posterior apex of abdomen, with no unusual features; each of its 6 setae nearly twice as long as greatest diameter of ring itself. Venter with multilocular disk pores not especially numerous on abdomen, occurring in irregular single rows along posterior and anterior borders across segments 8-5, very few along anterior margin of segment 4, in area posterior to vulva, absent elsewhere on body. Trilocular pores rather sparse, especially in certain "clear" areas on thorax. Oral-collar tubular ducts of same type and size as those on dorsum fairly numerous on abdominal segments, scattered on thorax and head. Body setae slender, some noticeably longer than those on dorsum. Circulus absent. Legs well developed, rather slender; hind coxae with very faint trace translucent pores, or these pores completely absent; hind tibiae slender, very small inconspicuous cluster translucent pores near apical part of segment, or these pores absent. Antennae normally 8-segmented, infrequently 7-segmented and this usually only on 1 side of body. Mouthparts comparatively broad. Notes. This species is rather closely related to Trionymus myersi McKenzie, but differs in having hind femora and tibiae quite slender and with, at most, only a faint trace of inconspicuous translucent pores near apical half of tibiae, • absent on femora; antennae normally 8-segmented; dorsal oral-collar tubular ducts small, all approximately the same size. In T. myersi, on the other hand, the hind femora and tibiae are robust, both provided with numerous, conspicuous translucent pores situated over most of segment; antennae normally 9-segmented, rarely 8segmented; and a few comparatively large dorsal oralcollar tubular ducts are associated with numerous smaller ducts of same general type. California Records (Map 185) Inyo Co.: 15 miles north Bishop, VII-17-63, on Elymus triticoides (D. R. Miller, UCD).

Trionymus haancheni McKenzie Haanchen Barley Mealybug (Figure 198) Trionymus haancheni McKenzie, 1960. Hilgardia 29(15) :765-768, illus. Type Locality and Host. Tulelake, Siskiyou County, California, on Haanchen barley, Hordeum vulgare (Gramineae). North American Distribution. Known only from California. Additional Hosts. Elymus sp. (Gramineae). External Features and Habitat. No information is available in external appearance of this mealybug. According to Osborn (1951), this species is found along the stems under the leaf sheaths of its host.

Map 185. Distribution in California of Trionymus

McKenzie.

furvus

Cottony egg clusters and accumulations of thick, sticky honeydew secretions are produced by this mealybug. Recognition Characters. Adult females, mounted, 2.00 to 5.10 mm long, 1.00 to 1.80 mm wide; body form, elongate, slender, more or less parallel-sided. Dorsum with cerarii present only on last 2 abdominal segments. Anal lobe cerarius with 2 moderately stout, conical setae, several slender auxiliary setae of various lengths, accompanied by scattered trilocular pores, all contained within oval area normally quite well sclerotized. Penultimate cerarius with 2 slightly smaller, conical setae, 3-4 slender auxiliary setae, small cluster trilocular pores, without basal sclerotization; no indication of cerarii appears anterior to these, even on head. Trilocular pores rather evenly beset on dorsum. Multilocular disk pores scattered on posterior 3 or 4 abdominal segments. Oral-collar tubular ducts of 2 sizes on abdomen, thorax, and head. Body setae narrow and slender. Anal ring apical, normal in form and size for genus; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores in considerable numbers in midregion of abdomen, few reaching lateral margin, from posterior apical forward to sixth segment. Trilocular pores rather numerous, generally distributed over venter. Oral-collar tubular ducts of 2 sizes, same as those on dorsum, larger ones more numerous, scattered over entire venter, although perhaps more numerous on abdomen. Body setae slender, of various lengths, some slightly longer than those on dorsum. Circulus rather small, quadrate-shaped, and, in most specimens, very definitely crossing intersegmental line between which it lies. Legs moderately slender in relation to length; hind coxa with translucent pores at base; claw without denticle on plantar

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Fig. 198. Trionymus haanchetii McKenzie, collected on Haanchen barley, vulgare (Gramineae), at Tulelake, Siskiyou County, California.

Hordeum

surface. Antennae normally 8-segmented, slender. Mouthparts comparatively short, stout, and blunt. Notes. This species was included on the type slides of Dysmicoccus modocensis Ferris ( = Trionymus modocensis [Ferris]), and was entangled with it until its description. It is, however, very evidently distinct from T. modocensis in possessing but 2 pairs of cerarii; presence of 30 or more dorsal multilocular disk pores on last 3 or 4 abdominal segments; and a comparatively short, stout, and blunt-type beak. In contrast, specimens of T. modocensis show a strong tendency toward the development of an antepenultimate pair of cerarii, this usually present, but may not be developed; 6 or less dorsal multilocular disk pores; and a definitely longer, acute-shaped beak (see Figure 198). H. T. Osborn (1951) reports this mealybug as of some economic concern: "On August 15th, the Siskiyou County Agricultural Commissioner's office at Tulelake submitted specimens of a mealybug reported to be so abundant in Haanchen barley in some fields that combines were being gummed up and clogged at harvest. When visited early in September, harvesting had been completed in the infested fields but large numbers of dead and dried-up mealybugs were present on the stubble. Large numbers of cottony egg clusters contained unhatched eggs. Broken straws on the ground and some standing uncut at the edge of the field showed evidence of feeding by mealybugs along the stems under the leaf sheaths. Accumulations of thick, sticky honeydew secretions were still present. Approximately 15,000 acres of Haanchen barley are planted in this district located south and west of Tulelake. Gumming of combines at harvest was said to have involved approximately 350 acres." California Records (Map 186) Siskiyou Co.: Tulelake, IX-4-51, on Hordeum vulgare (H. T. Osborn and W. E. Huse, C D A ) ; VIII-15-51, on Hordeum vulgare (culms) (W. E. Huse, C D A ) ; VI-15-54, on Hordeum vulgare (J. Schuh, USNM.) Ventura Co.: Ojai, 11 VII-22-59, on Elymus sp. (H. L. Wilson, C D A ) .

Trionymus mocus Ferris Mocker Grass Mealybug (Figure 199) Trionymus mocus Ferris, 1953. Atlas of scale insects of North America, Vol. VI: 490-491, illus. Type Locality and Host. San Mateo, San Mateo County, Cailfornia, on bamboo (undetermined species) (Gramineae). North American Distribution. Known only from California. Additional Hosts. To date known only from bamboo. External Features and Habitat. No available information on external appearance of this mealybug. 11 Specimens collected from this location under leaf sheaths of Elymus sp. have been tentatively identified as this species. Additional study material is needed, however, to positively ascertain the true identity of this mealybug.

Recognition Characters. Adult females, mounted, 1.50 to 2.50 mm long, 0.50 to 1.00 mm wide; body form slender, elongate oval. Dorsum with definite cerarii on last 2 abdominal segments. Anal lobe cerarius with 2 small conical setae, 1 or 2 slender auxiliary setae, little concentration of trilocular pores, no basal sclerotization. Penultimate cerarius with 2 small conical setae about which there is but few trilocular pores, no sclerotization. On abdominal segments 7 - 5 on each side near margin a pair of quite large, slender setae which may represent cerarii, these setae quite widely separated from each other. Trilocular pores rather sparsely beset on dorsum. Multilocular disk pores in a transverse row along both anterior and posterior borders of last 2 abdominal segments, along posterior border of segments 7-5, occurring singly over other parts of dorsum forward to head. Oral-collar tubular ducts very small, varying somewhat in size, either slightly larger or slightly smaller in diameter than a trilocular pore and but slightly longer than wide, present in extremely small numbers especially on abdominal segments, scattered on thorax and head. Body setae slender, variable in size, sparsely distributed on dorsum. Anal ring apical, presenting no distinctive features; each of its 6 setae about twice as long as diameter of ring itself. Venter with multilocular disk pores moderately numerous on abdomen in midregion from posterior apical segment forward to fifth segment, extending into lateral areas as far forward as head, scattered about singly elsewhere. Trilocular pores rather sparsely distributed, as on dorsum. Oral-collar tubular ducts similar to those on dorsum, scattered about sparsely, somewhat more abundant on abdominal segments 8 forward to 5. Body setae similar in size and distribution to those on dorsum.

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Fig. 199. Trionymus mocus Ferris, collected on bamboo (Gramineae), at San Mateo, San Mateo County, California.

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479

Circulus rather small, oval or round. Legs moderately slender in relation to length; hind coxae with cluster of translucent pores at base; claw without denticle on plantar surface. Antennae normally 7segmented, proportionately about same length as in forms having antennae 8-segmented. Notes. This species is rather closely related to Trionymus dolus Ferris but differs in the following characters: antennae 7-segmented; ventral multilocular disk pores present in lateral areas of anterior-most abdominal segments, thorax, and head; and anal lobe cerarius without surrounding sclerotized area. Trionymus dolus, on the other hand, possesses 8-segmented antennae; ventral multilocular disk pores are absent in lateral areas of anterior-most abdominal segments, thorax, and head; and anal lobe cerarius is surrounded by distinct area of sclerotization. California Records (Map 187) San Mateo Co.: San Mateo, V-27-52, on "bamboo" (E. L. Labadie Jr., C D A ) .

North American Distribution. Known only from California. Additional Hosts. Elymus condensatus and Elymus sp. (Gramineae). External Features and Habitat. No available information, except it is presumed to feed under leaf sheaths of its host. Recognition Characters. Adult females, mounted, 2.60 to 5.30 mm long, 1.00 to 2.50 mm wide; body form elongate, sides almost parallel. Dorsum with cerarii on last 3 abdominal segments, and infrequently frontal cerarius indicated on head. Anal lobe cerarius consisting of 2 stout, conical setae, surrounded by several quite large, slender, auxiliary setae, numerous but not crowded trilocular pores, all borne in quite large area of sclerotization. Penultimate cerarius with 2 smaller conical setae, no auxiliary setae, few associated trilocular pores. Antepenultimate cerarius consisting of 2 long, slender setae, 1 or 2 smaller, slender, auxiliary setae, small cluster trilocular pores. Frontal cerarius, when present, usually smiliar to antepenultimate cerarius, at times with cerarian setae appearing more conical. Trilocular pores fairly numerous, evenly beset on dorsum. Minute circular pores scattered. Oral-collar tubular ducts small, orifices not as great in diameter as trilocular pores, very numerous on abdomen, less abundant on thorax, almost disappearing in anterior end of body. Body setae slender, rather numerous, variable in length. Anal ring apical, normal for genus; each of its 6 setae about twice as long as diameter of ring itself. Venter with multilocular disk pores rather abundant in region posterior to vulva, along anterior and posterior borders of midregion of segments 8 and 7, very few on segment 6, absent anterior to this and elsewhere on body. Trilocular pores less numerous than on dorsum. Minute circular pores scattered. Oralcollar tubular ducts, of same type as those on dorsum, situated on posterior abdominal segments in numbers about equal to numbers of trilocular pores, less abundant on anterior abdominal segments, almost disappearing on thorax and head. Body setae similar in shape and size to those on dorsum, possibly slightly less abundant. Circulus quite small, quadrate-shaped, normally divided by segmental line of segments between which it lies. Legs well developed; translucent pores absent on segments; claw without denticle on plantar surface. Antennae 8-segmented, slender.

Trionymus modocensis (Ferris) Modoc Grass Mealybug

Notes. This species is very closely related to Trionymus haancheni McKenzie, and part of the type slides of T. modocensis included that species. Trionymus modocensis differs from T. haancheni mainly in the absence of dorsal multilocular disk pores on last 3 or 4 abdominal segments, these structures present in this area in the species last mentioned. Specimens collected at Camarillo, Ventura County, California, on Elymus condensatus, show slight structural divergences from the type for this species, such as 1 or 2 multilocular disk pores on dorsum of abdo-

(Figure 200) Dysmicoccus modocensis Ferris, 1953. Atlas of scale insects of North America, Vol. VI: 339-340, illus. ( i n p a r t ) . Trionymus modocensis (Ferris), McKenzie, 1960. Hilgardia 2 9 ( 1 5 ) : 7 6 4 and 767 (under notes). Type Locality and Host. One mile southeast of Alturas, Modoc County, California, on wild rye, Elymus triticoides (Gramineae).

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Fig. 200. Trionymus modocensis (Ferris), collected on wild rye, Elymus (Gramineae), southeast of Alturas, Modoc County, California.

triticoides

TAXONOMY men and more numerous multiloculars on venter of sixth abdominal segment. These differences appear to fall well within the normal range of variation for the species. California Records (Map 188) Lassen Co.: Susanville, IX-7-51, on Elymus triticoides (H. T. Osborn, CD A). Modoc Co.: 1 mile southeast of Alturas, IX-5-51, on Elymus triticoides (H. T. Osborn and L. White, C D A ) ; V I M 1-52, on Elymus triticoides (H. T. Osborn and L. White, C D A ) . Orange Co.: San demente, VIII-18-65, on Elymus ? sp. (G. Harper, C D A ) . Santa Barbara Co.: Montecito, IX-17-55, on "tall grass" (leaves) (G. G. Beevor, C D A ) . Siskiyou Co.: Tulelake, VII-10-52, on Elymus triticoides (W. Huse and H. T. Osborn, C D A ) ; IX4-51, on Elymus triticoides (H. T. Osborn and J. McKinney, C D A ) . Ventura Co.: Camarillo, VII-31-62, on Elymus condensatus (G. G. Beevor and D. Hare, C D A ) ; VIII-2-62, on Elymus sp. (G. G. Beevor and D. Hare, C D A ) .

Trionymus my ersi McKenzie Myers Mealybug (Figure 201) Trionymus my ersi McKenzie, 1961. Hilgardia 3 1 ( 2 ) : 48-51, illus. Type Locality and Host. Glendale, Los Angeles County, California, on Haworthia sp. (Liliaceae). North American Distribution. Known only from California.

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Additional Hosts. Aloe sp. (Liliaceae). External Features and Habitat. Unfortunately, no information is available on external appearance of this mealybug. According to the collector, the species prefers the leaf portion of the plant and does not go below ground onto the roots. Recognition Characters. Adult females, mounted, 1.60 to 2.60 mm long, 0.90 to 1.50 mm wide; body form somewhat elongate to rather broadly oval. Dorsum with cerarian pairs reduced, not more than 2 recognizable pairs on last 2 abdominal segments. Anal lobe cerarius with 2 moderately stout, conical setae, several slender auxiliary setae of various lengths, scattered trilocular pores, slight basal sclerotization; penultimate cerarius with 2 cerarian setae slightly smaller than those on anal lobes, several auxiliary setae, slight concentration trilocular pores, no sclerotization at base. Trilocular pores fairly numerous on dorsum, sparse in certain areas. Minute circular pores scattered. Multilocular disk pores present or absent; if present, very few, scattered over dorsum, these pores observed in only 2 of 22 specimens examined. Oral-collar tubular ducts of 3 distinct sizes, numerous, occurring on each abdominal segment, except last, and on thoracic segments and head. Anal ring apical, normal in size and form for genus; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores fairly numerous, occurring in area posterior to vulva, in an irregular single or double row along posterior borders of abdominal segments 8 to 5, few on anterior border of segments 8 to 7, occasional pore in lateral areas, absent on thorax and head, except in 1 specimen a single pore noted near clypeus. Trilocular pores sparsely beset on venter. Minute circular pores scattered. Oral-collar tubular ducts of 2 sizes present, larger ones more numerous, scattered over entire venter, perhaps more abundant on abdomen. Body setae slender, of various lengths, several noticeably longer than those on dorsum. Circulus absent. Legs rather robust; hind femora and tibiae with small translucent pores; denticle absent on plantar surface of claw. Antennae normally 9-segmented counting what appears to be a definite division of apical segment, although at times only 8 well developed segments present. Mouthparts fairly broad. Notes. This species is rather closely related to Trionymus smithii (Essig), but differs in possessing 2 distinct pairs of abdominal cerarii, and usually a 9segmented antenna, as compared to only an anal lobe pair of cerarii, and a 6- or 7-segmented antenna in the species last mentioned. Trionymus myersi also differs in having 3 different sizes of dorsal oral-collar tubular ducts as compared to only 2 sizes of these structures in T. smithii. According to correspondence from L. E. Myers, many of the infested Haworthia plants examined came from Germany and South Africa, which may or may

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Fig. 201. Trionymus myersi McKenzie, collected on Haworthia sp. (Liliaceae), at Glendale, Los Angeles County, California.

TAXONOMY not have been the origin of the mealybug. He reports that some of the varieties were quite heavily infested. California Records (Map 189) Los Angeles Co. : Glendale, V-24-45, on Haworthia sp. (M. Wagner, CDA); 1-7-46, on Haworthia sp. (M. Wagner, CDA); VII-17^6, on Aloe sp. (M. Wagner, LEM); Los Angeles, 11-26-45, on Haworthia sp. (F. Bacon and M. Wagner, CDA) ; Pacoima, 1-27-43, on Haworthia sp. (L. E. Myers, LEM); Pasadena, V-20-44, on Haworthia sp. (M. Wagner, CDA); XII-6-44, on Haworthia sp. (F. Bacon, LEM); Pomona, IV-23-41, on Haworthia sp. (J. H. Mitchell, CDA); V-10-44, on Haworthia sp. (L. E. Myers and J. B. Steinweden, LEM); XI-17-47, on Haworthia sp. (W. D. Dyer, CDA); West Los Angeles, V-27-43, on Haworthia sp. (Shilling, CDA). Merced Co.: Delhi, XI-24-43, on Aloe sp. (J. W. Dixon, UCD). San Diego Co.: Leucadia, VIII-8-55, on Haworthia sp. (D. Sprague and H. Wilson, CDA). Siskiyou

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Trionymus quadricircuius McKenzie, new species Square-Circulus Mealybug (Figure 202) Type Locality and Host. Type and paratype adult females on grass, Sitanion hystrix (roots) (Gramineae), 20 miles north of Bishop, Mono County, California, collected July 17, 1963, by D. R. Miller. Additional paratype adult females are available as follows, collected in California on species of Gramineae: on Sitanion hystrix (roots), Monitor Pass, Alpline County, collected August 2, 1964, by D. R.

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Miller and J. A. Froebe; and on unidentified grass, in Mountains west of Elsinore, Riverside County, collected October 22, 1938, by R. C. Dickson. Type Material. Holotype adult female (single specimen on slide) and female paratypes deposited in museum, University of California, Davis. North American Distribution. Known only from California. Additional Hosts. Unidentified grass (Gramineae). External Features and Habitat. This mealybug is rather small in size, is very lightly dusted with a white, powdery, wax exudation, and has a pinkish body color. Filamentation was not observed probably because of the small size of the mealybug. It is suspected, however, that at least 2 pairs of anal wax filaments would be present because of pore arrangement on derm. This species is found on the roots of grass, and careful examination failed to reveal its presence within the leaf sheath area. Recognition Characters. Adult females, mounted, 1.85 to 2.60 mm long, 0.95 to 1.50 mm wide; body form elongate, broadly oval, or rotund. Dorsum with cerarii present only on last 2 abdominal segments. Anal lobe cerarius consisting of 2 rather slender, conical setae, 4-5 slender auxiliary setae of various lengths, slight concentration trilocular pores, no basal sclerotization. Penultimate cerarius with 2 slightly smaller setae of same type, no auxiliary setae, slight concentration trilocular pores, no basal sclerotization. Trilocular pores evenly beset over entire dorsum. Multilocular disk pores in fairly uniform transverse band, or bands, across each abdominal segment, very sparsely scattered on prothorax. Oral-collar tubular ducts each about twice as long as wide with orifice about as great in diameter as trilocular pore, in irregular rows across abdominal segments, scattered over thorax and head. Body setae small, slender, fairly evenly beset on dorsum. Anal ring apical, without unusual characters; each of its 6 setae nearly twice as long as greatest diameter of ring itself. Venter with multilocular disk pores numerous, present on all abdominal segments, extending across segments to lateral margins, few scattered over thorax and head. Trilocular pores distributed rather evenly over venter. Oral-collar tubular ducts numerous especially on abdomen, fairly numerous and scattered on thorax and head. Body setae slender, approximately same length as those on dorsum. Circulus comparatively large, basically quadrateshaped (hence the specific name, quadricirculus), divided by intersegmental line between which it lies. Legs well developed; hind coxae with small translucent pores; claw without denticle on plantar surface. Antennae normally 8-segmented, infrequently 7-segmented and this usually only on 1 side of body. Mouthparts short and moderately broad. Notes This species associates most closely with Trionymus haancheni McKenzie, but differs mainly in the presence of multilocular disk pores in thoracic area on both body surfaces, but especially so on the

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Fig. 202. Trionymus quadricirculus McKenzie, new species, collected on grass, Sitanion hystrix (Gramineae), north of Bishop, Mono County, California.

TAXONOMY venter; and in an anal lobe cerarius without basal sclerotization. Trionymus haancheni, on the other hand, lacks multilocular disk pores in thoracic area on both body surfaces; and the anal lobe cerarius is surrounded by an oval area that is normally well sclerotized. California Records (Map 190) Alpine Co. : Monitor Pass, VIII-2-64, on Sitanion hystrix (roots) (D. R. Miller and J. A. Froebe, UCD). Mono Co.: 20 miles north Bishop, VII-17-63, on Sitanion hystrix (roots) (D. R. Miller, UCD). Riverside Co. : Mountains west of Elsinore, X-2238, on grass (R. C. Dickson, UCD).

Trionymus smithii (Essig) Smith Grass Mealybug (Figure 203) Ripersia smithii Essig, 1910. Pomona Col. Jour. Ent. 2(2) :218—221, illus, (in part). Pseudococcus smithii (Essig). Cockerell, 1916. Jour. Econ. Ent. 9:312-313, (in part). Trionymus smithii (Essig), Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., pp. 71-72, illus. Trionymus smithii (Essig), Ferris, 1950. Atlas of scale insects of North America, Ser. V: 273-274, illus. Type Locality and Host. Santa Paula, Ventura County, California, on wild rye, Elymus condensatus (Gramineae). North American Distribution. California and Mexico. Additional Hosts. Agrostis sp. prob, rubra (Gram-

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485

ineae), Chaetochloa caudata (Gramineae), Elymus triticoides, Elymus spp. (Gramineae), Secale cereale (Gramineae), and undetermined grasses (Gramineae). External Features and Habitat. The adult female of this mealybug has a body that is decidedly long and narrow, or elongate-elliptical in form. There may be as many as 3 pairs of thin caudal filaments, though these are rudimentary and often imperfect. The body is pinkish to slate-colored, and is lightly covered with a fine, white, waxy secretion that scarcely hides the body color, with greatest percentage of this secretion produced near posterior apex of body. The legs are white. No true ovisac has been found associated with this mealybug, and it is suspected that the species is ovoviviparous, since first instar nymphs are commonly found hidden underneath the abdomen of the adult female. When disturbed, a whitish honeydew is expelled through the ostioles. This species occurs beneath the leaf blade sheaths of its host. Recognition Characters. Adult females, mounted, 2.00 to 5.50 mm long, 0.90 to 2.00 mm wide; body form elongate with nearly parallel sides. Dorsum with cerarii present only on anal lobes. Anal lobe cerarius with 2 rather small, conical setae, no closely accompanying slender auxiliary setae, either no concentration trilocular pores or with very small group of less than 10 immediately about bases of setae. Trilocular pores evenly beset on dorsum. Minute circular pores numerous, scattered over dorsal surface. Multilocular disk pores very few, situated on last 2 or 3 abdominal segments, and near lateral margins of segments 7 forward to 5. Oral-collar tubular ducts small, orifices with diameter equaling that of a trilocular pore, slightly longer than wide, few scattered on last 3 or 4 abdominal segments. Body setae very few, small and slender. Anal ring of normal form and size for genus; its 6 setae slightly longer than greatest diameter of ring. Venter with multilocular disk pores relatively few, situated in midregion of posterior apical segment of abdomen forward to sixth, or perhaps fifth segment, also in small numbers in lateral areas, few such pores usually present on head anterior to clypeus. Trilocular pores generally distributed on venter. Minute circular pores scattered. Oral-collar tubular ducts of same size and type as those on dorsum occur especially in lateral areas of abdominal segments, also in limited numbers in midregion of segments 9 forward to 5, very few such ducts occur along submargin of thorax, very few situated on head between antennal bases. A few exceedingly minute, oral-collar type tubular ducts occur in midregion of abdomen. Body setae sparse, slender, variable in length, some longer than those on dorsum. Circulus absent. Legs well developed, comparatively small; hind coxae definitely larger than those of other legs, thickly beset with conspicuous translucent pores; claw without denticle on plantar surface. An-

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Fig. 203. Trionymus smithii (Essig), collected on wild rye, Elymus condensatus (Gramineae), at Santa Paula, Ventura County, California.

tennae normally 7-segmented, at times varying from 6- to even 8-segmented. Notes. The identification of this species is based upon specimens from near type locality which were compared with the types by Ferris many years ago. It was noted at that time (1918) that the type slide material included 2 species, and the name smithii was definitely restricted by Ferris to one of these. Ferris states: "There is some variation in the specimens at hand. In some the conical setae of the anal lobe cerarii are slightly larger than here illustrated and are accompanied by a small cluster of pores. In some the posterior coxae are not as thickly beset with pores as is here illustrated, although they are similarly enlarged. In some cases there are perhaps a few more multilocular pores. But on the whole all the available specimens seem to come within what are probably normal limits of variation." This species is closely related to Trionymus myersi McKenzie, but differs in possessing only an anal lobe pair of cerarii, and very few dorsal, short oralcollar tubular ducts restricted to last 3 or 4 abdominal segments, whereas in T. myersi the cerarii are developed on last 2 abdominal segments,and there are rather numerous, small dorsal oral-collar tubular ducts of 3 different sizes situated on all abdominal segments, except the last, and on thorax and head. Also the hosts attacked are quite different for the species, since T. smithii seems restricted to members of the Gramineae, while T. myersi has been taken only on representatives of the Liliaceae. California Records (Map 191) Inyo Co.: 3 miles south Bishop, VIII-19-63, on Gramineae (leaf sheath) (D. R. Miller and J. A. Froebe, U C D ) . Los Angeles Co.: Whittier (Puente Hills), 1921, on "broom-like grass" (H. M. Armitage, C D A ) . Modoc Co.: 4 miles west Alturas, VI-27-63, on Secale cereale (grass sheath) (D. R. Miller, U C D ) . Riverside Co.: 3.4 miles northeast Gilman Springs, III-9-63, on Agrostis sp. probably rubra (D. R. Miller, U C D ) . San Benito Co.: San Juan Grade, XII-1-17, on Elymus sp. (G. F. Ferris, U C D ) . Santa Barbara Co.: Cuyama, IV-29-55, on "native grass" (G. G. Beevor, C D A ) . Santa Clara Co.: Santa Clara Valley, XII-20-16, on "grass" (R. W. Doane, C D A ) ; Stanford University, 1-6-18, on "perennial grass" (G. F. Ferris, UCD). Santa Cruz Co.: Santa Cruz, XI-20-45, on "lawn grass" (H. Washburn, CAS). Ventura Co.: Camarillo, VII-31-62, on Elymus condensatus (G. G. Beevor and D. Hare, C D A ) ; Ojai, VII-22-59, on Elymus sp. (under leaf sheath) (H. L. Wilson, C D A ) ; Santa Paula, VII-1910, on Elymus condensatus (P. E. Smith, CAS); Springville, VI-11-47, on rye grass (perennial) (P. B. Travis, CDA). Yolo Co.: Plainsfield, VIII-24-63, on Elymus triticoides (grass sheath) (D. R. Miller, U C D ) .

Map 191. Distribution in California of Trionymus smithii (Essig). M*rin Sa/% frsneiteo Sen Matte Santa Cruz

Trionymus utahensis (Cockerell) Utah Grass Mealybug (Figure 204) Pseudococcus neomexicanus var. utahensis Cockerell, 1916. Jour. Econ. Ent. 9:313. Trionymus bromi Ferris, 1918. Stanford Univ. Pubs., Univ. Ser., p. 67. Trionymus bromi Ferris, 1920. Stanford Univ. Pubs., Univ. Ser., Biol. Sci. 1 ( 1 ) : 3 1 . Trionymus bromi Ferris, 1950. Atlas of scale insects of North America, Ser. V:255-256, illus. Trionymus utahensis (Cockerell), McKenzie and Williams, 1965. Hilgardia 3 7 ( 1 ) : 11-14, illus. Type Locality and Host. Salt Lake City, Salt Lake County, Utah, on wild rye, Elymus sp. (Gramineae). North American Distribution. California, Oregon, and Utah. Additional Hosts. Ammophila arenaria (Gramineae), Avena barbata (Gramineae), Bromus sp. prob. orcuttianus (Gramineae), Bromus sp. (Gramineae), Cynodon dactylon (Gramineae), Ericameria fasciculata (Compositae), Sorghum halepense (Gramineae), and Triticum aestivum (Gramineae). External Features and Habitat. Cockerell (1916) states the following about external appearance of this mealybug in the original description: "Apparently when the females reach a fairly large size (about half grown) they become enclosed in rather thin cottony sacs of rather close texture. A female that was removed from its situs and placed in a vial, soon entirely enclosed itself in a cylindrical cottony mass about 6 mm long and 2 mm diameter. This sac was apparently much more abundantly developed and certainly considerably more fluffy than in specimens taken from the stems. Those in stems were of course

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Fig. 204. Trionymus utahensis (Cockerell), collected on Bermuda grass, Cynodon dactylon (Gramineae), at Sacramento, Sacramento County, California. (Illustration prepared from specimens collected on Bromus sp. [Gramineae], near Stanford University, Santa Clara County, California.)

T AXON

much flattened and the cottony secretion pressed together. A female with egg-mass of 103 eggs, collected on Nov. 23, was found at one end of sac, and not covered up, at least when leafsheath was torn away; neither were the eggs entirely covered up, but the cottony fibers being sticky (but not viscid) tenaciously held the eggs in place. The body coloration was not noted, certainly not striking, and if I remember right pale yellowish. The females removed from their sacs were covered with a thin white meal, and nearly destitute of the usual lateral and caudal secretions. The exhausted female found on Nov. 23 turned dull, dark, crimson-brown when boiled in KOH. The egg was described as pale yellowish-brown, darker at one end, about .4 mm long by .2 mm diameter." This mealybug is found concealed between the sheaths and stems of grass, and hence cannot be observed in an undisturbed condition. Recognition Characters. Adult females, mounted, 3.90 to 5.00 mm long, 1.30 to 1.95 mm wide; body form elongate with nearly parallel sides. Dorsum with cerarii present only on anal lobes, those with 2 rather slender, conical setae, 2-3 accompanying slender auxiliary setae, very slight concentration trilocular pores, no basal sclerotization. Trilocular pores evenly beset on dorsum. Multilocular disk pores rather few, situated in transverse bands across abdomen from apical segment forward to at times fifth segment, absent elsewhere on dorsum, except infrequently 1-4 such pores scattered on thorax. Oral-collar tubular ducts of 2 slightly different sizes on abdomen mostly in irregular rows along posterior border of each segment, along lateral margin from abdomen to head, sparsely scattered over thoracic areas. The orifice diameter of these ducts is somewhat greater than that of a trilocular pore, their length perhaps 2-3 times their diameter. Body setae small and slender. Anal ring apical, with no unusual features; each of its 6 setae about twice as long as greatest diameter of ring. Venter with multilocular disk pores situated on posterior apical segment of abdomen forward to fourth segment, mostly in an irregular single or double row along posterior border of midregion, along lateral margins as far forward as head, very few scattered on thorax. Trilocular pores fairly evenly beset on venter. Oral-collar tubular ducts of 2 sizes, similar to those on dorsum, present in considerable numbers in lateral regions from ninth abdominal segment to head, more sparsely distributed especially in midregion of abdomen and thorax. Body setae slender, sparsely distributed on venter. Circulus quite small, circular or oval, not divided by segmental fold. Legs quite small, not especially stout; hind coxae with cluster translucent pores at base; claw without denticle on plantar surface. Antennae 8-segmented. Notes. McKenzie and Williams (1965) determined that Trionymus bromi Ferris is definitely a synonym of T. utahensis (Cockerell). Trionymus utahensis has several dorsal multilocular disk pores on the fifth abdominal segment, and its thorax has more sub-

OMY

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marginal multiloculars on the venter than illustrated for T. bromi by Ferris. Also the circulus is slightly larger, and the translucent pores on the hind coxae are more abundant than shown in his drawing. Ferris mentioned a few of these deviations, especially in examples collected on wheat near Salt Lake City, Utah. Trionymus utahensis is very closely related to Trionymus caricis McConnell, but differs in having the multilocular disk pores of dorsum confined to the last 5 abdominal segments; an 8-segmented antenna; and longer oral-collar tubular ducts. In the species last mentioned, the multiloculars are scattered over entire dorsum; the antenna is normally 7-segmented (at times even 6-segmented); and the oral-collar tubular ducts are noticeably shorter. California Records (Map 192) Los Angeles Co.: Pasadena, IV-10-16, on Sorghum halepense (A. G. Smith, CAS). Monterey Co.: Pacific Grove, XII-1-17, on Ericameria fasciculata (G. F. Ferris, UCD). Napa Co.: Napa, VII-2-64, on grass (H. L. McKenzie and D. R. Miller, UCD). Placer Co.: 3 miles west Applegate, VIII-8-64, on Gramineae (leaf sheaths) (D. R. Miller, UCD). Plumas Co.: Lake Almanor, IX-6-51, on Bromus sp. prob. orcuttianus (H. T. Osborn, CDA). Sacramento Co.: Sacramento, 1918, on Cynodon dactylon (H. S. Smith, UCD). San Francisco Co.: San Francisco (sand dunes), 1-10-18, on Ammophila arenaria (G. F. Ferris, UCD). Santa Clara Co.: Near Congress Springs, X-15-16, on Bromus sp. (G. F. Ferris, UCD); near Stanford University (Corte Madera Creek), X-3-16, on Bromus sp. (P. W. Doane, UCD). Sierra Co.: Gold Lake, IX-15-65, on grass (T. R. Haig, CDA). Tehama Co.: Yolla Bolly, Middle Eel, X-5-51, on Bromus sp. (H. T. Osborn, CDA).

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Trionymus

winnemucae McKenzie, new species Winnemuca Grass Mealybug (Figure 205)

Type Locality and Host. Winnemuca Lake, Alpine County, California, on presumably wheat grass, Agropyron ? sp. (Gramineae), collected June 14, 1964, by J. A. Froebe. Type Material. Holotype adult female (single specimen on slide) and paratypes in museum, University of California, Davis. North American Distribution. Known only from California. Additional Hosts. None External Features and Habitat. No available information relative to external appearance of this species. The mealybug occurs in the leaf-blade sheaths of its grass host. Recognition Characters. Adult females, mounted, 2.00 to 3.15 mm long, 0.95 to 1.30 mm wide; body form elongate, slender, with almost parallel sides. Dorsum with cerarii present only on anal lobes. Anal lobe cerarius with 2 rather small, conical setae, no closely associated slender auxiliary setae, slight concentration trilocular pores, no surrounding sclerotization. Trilocular pores evenly beset on dorsum. Minute circular pores scattered. Multilocular disk pores rather sparse, scattered on last 5 or 6 abdominal segments, perhaps more abundant on eighth and seventh segments, very few in head region. Tubular ducts absent on dorsum. Body setae small and slender, sparsely scattered over dorsal surface, certain "clear" areas evident especially on thorax. Anal ring apical, normal in form and size for members of this genus; each of its 6 setae only slightly longer than greatest diameter of ring. Venter with multilocular disk pores comparatively few, situated in both midregion and lateral areas from posterior apical segment of abdomen to segment 4, scattered along submargin of anterior abdominal segments, thorax, and head, clusters of these pores especially noticeable around spiracles and below eye, absent in sternal areas. Trilocular pores sparsely beset on venter. Minute circular pores scattered. Oral-collar tubular ducts very small, relatively few, scattered on most abdominal segments, perhaps more numerous on posterior half of abdomen, apparently absent on thorax and head. Body setae slender, of various lengths, some slightly longer than those on dorsum.

Circulus absent. Legs comparatively small; hind coxae equipped with rather large translucent pores; hind tarsus almost half as long as tibia; claw without denticle on plantar surface. Antennae 7-segmented. Mouthparts moderately broad. Notes. This species shares with Trionymus smithii (Essig) only an anal lobe pair of cerarii. It differs from T. smithii in having ventral multilocular disk pores below eye and near both pairs of spiracles; proportionately smaller hind coxae with fewer translucent pores; fewer trilocular pores near orifice of posterior spiracles; and apparently occurring at higher altitudes (approximately 8,000 feet). Trionymus smithii, on the other hand, has no ventral multilocular disk pores below eye or near thoracic spiracles; hind coxae definitely larger than those of other pairs, and thickly beset with translucent pores; trilocular pores numerous, clustered near orifice of posterior spiracles; and normally occurring near sea level. California Records (Map 193) Alpine Co.: Winnemuca Lake, VI-14-64, on Agropyron ? sp. (J. A. Froebe, U C D ) .

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OF

SPECIES

Fig. 205. Trionymus winnemucae McKenzie, new species, collected presumably on wheat grass, Agropyron? sp. (Gramineae), at Winnemuca Lake, Alpine County, California.

491

DISTRIBUTION TABLE

The table presents distributional information representing each pseudococcid species by counties in California. In addition, there is a column recording total number of species for each county. Although several counties have few mealybug species recorded for them, it is quite certain that representatives are present; and that if surveys and collections were conducted, mealybugs would be subsequently discovered.

efforts of the late Professor G. F. Ferris). In other instances, however, distribution and host records were made available through an actual examination of slide specimens represented in the National Collection of Coccoidea (U. S. National Museum), Washington, D. C.; the insect collections of the California State Department of Agriculture, Sacramento; the California Academy of Sciences, San Francisco; private collection of L. E. Myers, Los Angeles, California, and the University of California, Department of Entomology, collection at Riverside (this collection recently donated to the museum, University of California, Davis).

Information used to prepare this table was taken mainly from the extensive collections in the museum, University of California, Davis (formerly the Stanford University Collection, accumulated through the

493

Amonostherium lichtensioides Anisococcus abnormalis Anisococcus adenostomae Anisococcus crawii Anisococcus didymus Anisococcus ephedrae Anisococcus imperialis Anisococcus oregonensis Anisococcus quercus Anthelococcus simondsi Antonina crawii Antonina graminis Antonina pretiosa Cataenococcus olivaceus Chnaurococcus trifolii Chnaurococcus villosa Chorizococcus abortivus Chorizococcus abroniae Chorizococcus brevicrurus Chorizococcus californicus Chorizococcus coxindex Chorizococcus fistulosus Chorizococcus interruptus Chorizococcus lounsburyi Chorizococcus penultimatus Chorizococcus polyporus Chorizococcus psoraleae Chorizococcus reducta Chorizococcus rostellum Chorizococcus senarius Chorizococcus snellingi Chorizococcus wilkeyi Chorizococcus wilsoni Chorizococcus yuccae Crisicoccus azaleae Crisicoccus pini Cryptoripersia salina Cryptoripersia tubulata Cucullococcus vaccinii

494 DISTRIBUTION

Alameda Alpine Amador Butte Calaveras Colusa Contra Costa Del Norte El Dorado Fresno Glenn Humboldt Imperial Inyo Kern Kings Lake Lassen Los Angeles Madera Marin Mariposa Mendocino Merced Modoc Mono Monterey Napa Nevada Orange Placer Plumas Riverside Sacramento San Benito San Bernardino San Diego San Francisco San Joaquin San Luis Obispo San Mateo Santa Barbara Santa Clara Santa Cruz Shasta Sierra Siskiyou Solano Sonoma Stanislaus Sutter Tehama Trinity Tulare Tuolumne Ventura Yolo Yuba X

X X

X

X X X

X

X X

X

T

X

X X

ABLE

X X

X

X

X

X

X

X X XX X

X

X

X

X

X

X X

X

X

X

X X X

X X

XX X X X

X

X

X

X

X

X

X

X

X

X

X X

X X X X X

X X

X X

X

X X

X

X

X

X

X

X X X X X X

X X X X X

X X

X

X

X

X X

X X X

X

X X

X X

X

X

X X X X X

X

X

X

X X X

X

X

X

X

X

X X X

X X X

XXX X

X

X X X

X

X X

X X

X

X

X

X

Discococcus caricis Discococcus flabellatus Discococcus graminis Discococcus hopi Discococcus simplex Discococcus spectabilis Distichlicoccus arundinis Distichlicoccus californicus Distichlicoccus megacirculus Distichlicoccus salinus Dysmicoccus aciculus Dysmicoccus brevipes Dysmicoccus mackenziei Dysmicoccus pinicolus Dysmicoccus quercicolus Dysmicoccus racemus Dysmicoccus ryani Dysmicoccus timberlakei Dysmicoccus vacuatus Ehrhornia cupressi Ferrisia virgata Heliococcus adenostomae Heliococcus atriplicis Heliococcus stachyos Heterococcus arenae Heterococcus pulverarius Humococcus atriplicis Humococcus caritus Humococcus hilariae Humococcus inornatus Hypogeococcus spinosus Misericoccus arenarius Nipaecoccus aurilanatus Nipaecoccus nipae Paludicoccus distichlium Phenacoccus alleni Phenacoccus artemisiae Phenacoccus cajonensis Phenacoccus celtisifoliae

DISTRIBUTION TABLE

X X X Alameda X X X Alpine Amador Butte Calaveras Colusa Contra Costa Del Norte X El Dorado X X Fresno X X Glenn Humboldt X Imperial X X X X Inyo X X X X Kern X X X X X X Kings Lake Lassen X Los Angeles X X X X X X X X X X X Madera Marin Mariposa X X Mendocino X Merced X X Modoc X X X X X Mono X Monterey X X X Napa X X Nevada X X X X X Orange X Placer X Plumas X X X X X Riverside Sacramento San Benito X X San Bernardino X X X X X X X X X San Diego X X X X X X X X X X San Francisco X X San Joaquin X X X San Luis Obispo X X X San Mateo X X X Santa Barbara X Santa Clara X X X X X X X X Santa Cruz X X X Shasta Sierra X Siskiyou X X Solano X X Sonoma Stanislaus Sutter Tehama Trinity Tulare Tuolumne X X Ventura X Yolo X X X Yuba

495

X X X X X

X X X

X

X XXX

X X

X X X X

X

X

X X

Phenacoccus colemani Phenacoccus dearnessi Phenacoccus defectus Phenacoccus destitutus Phenacoccus dicoriae Phenacoccus echeveriae Phenacoccus eremicus Phenacoccus eriogoni Phenacoccus eschscholtziae Phenacoccus giganteus Phenacoccus gossypii Phenacoccus graminosus Phenacoccus helianthi Phenacoccus infernalis Phenacoccus lotearum Phenacoccus lycii Phenacoccus megaulus Phenacoccus minimus Phenacoccus multisetosus Phenacoccus pauperatus Phenacoccus solani Phenacoccus solenopsis Planococcus citri Planococcus kraunhiae Pseudantonina arundinariae Pseudococcus aberrans Pseudococcus dispar Pseudococcus diversus Pseudococcus fragilis Pseudococcus importatus Pseudococcus longisetosus Pseudococcus longispinus Pseudococcus maritimus Pseudococcus megasetosus Pseudococcus microcirculus Pseudococcus obscurus Pseudococcus pertusus Pseudococcus pseudobscurus Pseudococcus sorghiellus

496 DISTRIBUTION

Alameda Alpine Amador Butte Calaveras Colusa Contra Costa Del Norte El Dorado Fresno Glenn Humboldt Imperial Inyo Kern Kings Lake Lassen Los Angeles Madera Marin Mariposa Mendocino Merced Modoc Mono Monterey Napa Nevada Orange Placer Plumas Riverside Sacramento San Benito San Bernardino San Diego San Francisco San Joaquin San Luis Obispo San Mateo Santa Barbara Santa Clara Santa Cruz Shasta Sierra Siskiyou Solano Sonoma Stanislaus Sutter Tehama Trinity Tulare Tuolumne Ventura Yolo Yuba X

X

X X X

X X X

X X X

X

TABLE

X X X

X

X

X

X X

X X

X

X

X X X

X

X

X

X X X

X

X X X X X

X X X

X X X X

X X X X X X X X X X X X X X X

X

X X

X X X X X X XXX X X X X X X X X XXX

X X X X X X X X X

X

X

X

X X X X

X

X X

X

X

X X X X X

X X X X

X X X X X

XXX

X X X

X

X

X X

X X X X X X

X

X

X X X X X X X X X X X X X

X X X X X X X X X X X X X X X X X X X X X X

X

X

X

X

X

X X X X X

X

X X X

X X

X X X X

X X

X

X

X X X X X X X X X

X X

X

X

X

X

X

X X X X

XXX X

X X

X X

X

X

X

X X

X X X

X

X X X X X X X X

X X X X X

X X

X

X

X X

X X X X X X X X X X X

X X

X

X

X X XXX X

XX X

X

X X

XX X X X X X

X

X X

X XXX X X

X X X X X X

X X X X X

X X X

X

X

X X X X X

X X X

Pseudococcus sparsus Puto acirculus Puto albicans Puto ambiguus Puto arctostaphyli Puto atriplicis Puto bryanthi Puto californicus Puto cupressi Puto decorosus Puto echinatus Puto laticribellum Puto mimicus Puto nulliporus Puto pacificus Puto pricei Puto profusus Puto simmondsiae Puto yuccae Pygmaeococcus morrisoni Radicoccus kelloggi Rhizoecus associatus Rhizoecus bicirculus Rhizoecus bituberculatus Rhizoecus browni Rhizoecus cacticans Rhizoecus californicus Rhizoecus cyperalis Rhizoecus disjunctus Rhizoecus eluminatus Rhizoecus falcifer Rhizoecus gracilis Rhizoecus graminis Rhizoecus kondonis Rhizoecus leucosomus Rhizoecus pritchardi Rhizoecus sonomae Rhizoecus spinosus Scaptococcus californicus

DISTRIBUTION

Alameda Alpine Amador Butte Calaveras Colusa Contra Costa Del Norte El Dorado Fresno Glenn Humboldt Imperial Inyo Kern Kings Lake Lassen Los Angeles Madera Marin Mariposa Mendocino Merced Modoc Mono Monterey Napa Nevada X Orange Placer Plumas Riverside Sacramento San Benito San Bernardino San Diego San Francisco San Joaquin San Luis Obispo San Mateo Santa Barbara Santa Clara Santa Cruz Shasta Sierra Siskiyou Solano Sonoma Stanislaus Sutter Tehama Trinity Tulare Tuolumne Ventura X Yolo Yuba X X X

X

X

X

X X

X X X

X X

X X

XX

X X X

X X

X X X

X X X

X X

X

X

X

X

X X X

X X X X X X

X

X X

X X

X

X

X

X

X

X

X

X

TABLE

xx X X

X

X

X

X X

X X X

X X

X

X

X X X

X X

X

x~

X X

X X X

X

X X X

X

X X

X X

X XX X X

XXX X

X

X

497

X

X X X

X X

X

X X X

X X

X X

X

X X X

X X X X

X X

X X

XX X X X

XXX X X X

X

X X X

X

X

X

X

Scaptococcus milleri Spilococcus andersoni Spilococcus atriplicis Spilococcus cactearum Spilococcus ceanothi Spilococcus corticosus Spilococcus eriogoni Spilococcus geraniae Spilococcus gutierreziae Spilococcus implicatus Spilococcus keiferi Spilococcus larreae Spilococcus parkeri Spilococcus parvicirculus Spilococcus pressus Spilococcus prosopidis Spilococcus quercinus Spilococcus sequoiae Spilococcus ventralis Stemmatomerinx decorata Tridiscus distichlii Tridiscus multiorbis Trionymus americanus Trionymus caricis Trionymus diminutus Trionymus dolus Trionymus festucae Trionymus frontalis Trionymus furvus Trionymus haancheni Trionymus mocus Trionymus modocensis Trionymus myersi Trionymus quadricirculus Trionymus smithii Trionymus utahensis Trionymus winnemucae Total Number Mealybug Species

498 DISTRIBUTION TABLE

X Alameda X X X X X Alpine X Amador Butte Calaveras Colusa Contra Costa X X X Del Norte El Dorado Fresno X Glenn Humboldt X X X Imperial X X X X X X X Inyo X X Kern X X X X X X Kings Lake Lassen Los Angeles X X X .X X X X Madera Marin X X Mariposa Mendocino Merced X Modoc X X Mono X Monterey X X X Napa X X Nevada X X X Orange X X Placer Plumas X X Riverside X X X X X X Sacramento X X X San Benito San Bernardino X X X X X X San Diego X X X X X X X X X San Francisco X X X San Joaquin X X San Luis Obispo X X X San Mateo X X Santa Barbara X Santa Clara X X X X X Santa Cruz X X X X X Shasta X Sierra Siskiyou X X X X Solano X Sonoma X Stanislaus X X Sutter Tehama Trinity X X Tulare X Tuolumne X Ventura X X X X Yolo X Yuba X

X

X

X

X

X

X

X X

X X

X

X

X

X X XXX

X

X

X

X

X

X

X

X X

X X X

X

X X X X

X XX

X

X

X

X X X X

X

X

X XX X

X

X

X

37 11 0 13 1 3 14 5 8 17 4 13 29 29 25 2 4 9 59 2 17 4 9 7 18 6 21 9 13 34 11 12 56 18 6 50 50 19 13 9 28 19 45 21 6 8 15 13 11 10 2 7 7 11 4 29 19 6

LITERATURE CITED

(General references only. Specific taxonomic literature for each California mealybug is included under species concerned.) Borkhsenius, N. S. 1 1947. On the taxonomic significance of morphological characters of mealybugs (Coccoidea, Pseudococcus). Akad. Nauk. Dok. SSSR (Moscow) (n. s. 58) 9:2109, paragraph 3 (In Russian). 1948. Notes on Pseudococcus comstocki (Kuw.) and some allied species (Homoptera: Coccoidea), with descriptions of three new species. Bull. Ent. Res. 39(3) :417-421 (In English). 1949. Fauna of USSR. Homoptera, Pseudococcidae. Akad. Nauk. Zool. Inst. (Moscow) (n. s. 38) : 237-241, illus. (In Russian). Bouché, P. F. 1844. Beiträge zur naturgeschichte der scharlachlause (Coccina). Stettin. Ent. Ztg. 5:302. Brain, C. K. 1912. Contribution to the knowledge of mealybugs, genus Pseudococcus, in the vicinity of Cape Town, South Africa. Ann. Ent. Soc. Amer. 5 ( 2 ) : 177-189, illus. Carnes, E. K. 1907. The Coccidae of California. Second biennial report for 1905-1906. Calif. State Comnr. Hort., pp. 155-222, illus. DeLotto, G. 1958. The Pseudococcidae (Horn.: Coccoidea) described by C. K. Brain from South Africa. Bull. Brit. Mus. (Nat. Hist.) Ent. 7(3) :79120, illus. Essig, E. O. 1909. The genus Pseudococcus in California. Pomona Col. lour. Ent. 1:35-46, illus. 1914. The mealybugs of California. Calif. Comn. Hort. Monthly Bull. 3(3):97-143, illus. 1

1942.

College entomology. New York, Macmillan. 900 pp., illus. Ezzat, Y. M., and H. S. McConnell. 1956. The mealybug tribe Planococcini (Pseudococcidae, Homoptera). Md. Agr. Expt. Sta. Bull. A-84:1-108, illus. Fernald, IVI. E. 1903. A catalogue of the Coccidae of the world. Mass. Agr. Expt. Sta. Spec. Bull. 88. 360 pp. Ferris, G. F. 1918. The California species of mealybugs. Stanford Univ. Pubs. Univ. Ser. 78 pp., illus. 1950. Atlas of the scale insects of North America. Series V. The Pseudococcidae (Part I). Stanford University Press, Stanford, pp. 1-278, illus. 1953. Atlas of the scale insects of North America. Volume VI. The Pseudococcidae (Part II). Stanford University Press, Stanford, pp. 279506, illus. Goux, L. 1941. Contribution a l'etude d'un faisceau d'espèces constituant un sous-genre nouveau du genre Pseudococcus (Hem. Coccidae). Bull. Mus. d'Hist. Nat. Marseille l ( l ) : 6 6 - 8 3 , illus. Green, E. E. 1925. Observations on British Coccidae, IX. Ent. Monthly Mag. 61:40-41, illus. Hall, W. J. 1923. Further observations on the Coccidae of Egypt. Egypt Min. Agr. Tech, and Sci. Serv. Bull. 36:12-13, illus. Hambleton, E. J. 1946. Studies of hypogeic mealybugs. Rev. de Ent. 17(1-2): 1-77, illus. MacGillivray, A. D. 1921. The Coccidae. Urbana, 111. Scarab. 502 pp. McKenzie, Howard L. 1960. Taxonomic study of California mealybugs, with descriptions of new species (Homop-

According to British Museum System of Transliteration. 499

500

LITERATURE

CITED

tera: Coccoidea: Pseudococcidae). Hilgardia 29(15) :681-770, illus. 1961. Second taxonomic study of California mealybugs, with descriptions of new species (Homoptera: Coccoidea: Pseudococcidae). Hilgardia 31(2) :15-52, illus. 1962. Third taxonomic study of California mealybugs, including additional species from North and South America (Homoptera: Coccoidea: Pseudococcidae). Hilgardia 32( 14) :637-688, illus. 1964. Fourth taxonomic study of California mealybugs, with additional species from North America, South America, and Japan (Homoptera: Coccoidea: Pseudococcidae). Hilgardia 35( 10) :211-272, illus. McKenzie, Howard L. and D. J. Williams. 1965. Fifth taxonomic study of North American mealybugs, with revisionai notes on seven species (Homoptera: Coccoidea: Pseudococcidae). Hilgardia 3 7 ( 1 ) : 1-15, illus.

Morrison, H. 1945. The mealybug genus Heterococcus Ferris and some of its relatives (Homoptera: Coccoidea). Jour. Wash. Acad. Sci. 35(2):38-55, illus. Morrison, H., and Alice V. Renk. 1957. A selected bibliography of the Coccoidea. U. S. Dept. Agr. Misc. Pub. 734. 222 pp. Morrison, H., and E. R. Morrison. 1965. A selected bibliography of the Coccoidea. First supplement. U. S. Dept. Agr. Misc. Pub. 987. 44 pp. Takahashi, R. 1958. Key to the genera of Pseudococcidae in Japan, with descriptions of three new genera and two new species (Homoptera). Osaka Prefecture Univ. Bull. (ser. B) 7:1-8, illus. Williams, D. J. 1962. The British Pseudococcidae (Homoptera: Coccoidea). Bull. Brit. Mus. (Nat. Hist.) Ent. 12:1-79, illus. Zimmerman, E. C. 1948. Insects of Hawaii, Homoptera: Sternorhyncha. Univ. Hawaii Press, 5:1-464, illus.

HOST INDEX (With Associated Mealybugs) Both the common 1 and the scientific names of host plants are arranged alphabetically to assist in finding desired information. The associated mealybug species are recorded, according to their scientific names only, in alphabetical sequence either under a plant genus when several species of that host are involved, or under a single respective host species. In setting up the HOST INDEX, an attempt has been made to record every known host for each California mealybug species with the exception of those forms which are obviously general feeders. In these cases records are made of only a few of the more preferred hosts as indicated under the "Additional Hosts" section, as well as all host plants for specimens identified f r o m microscope slide mounts included in the "California Records" section.

Abies sp. (Pinaceae), 360 Puto cupressi, 343 Puto profusus, 360 Abies concolor (Pinaceae), 343 Abronia sp. (Nyctaginaceae), 93, 276 Chorizococcus abroniae, 91 Phenacoccus solani, 273 Puto echinatus, 347 Abronia latifolia (Nyctaginaceae), 91, 93 Abronia marítima (Nyctaginaceae), 91, 92, 93 Abronia umbellata (Nyctaginaceae), 347, 350 Abutilón sp. (Malvaceae), 253, 254, 389 Phenacoccus gossypii, 248 Rhizoecus falcifer, 389 Acacia sp. (Leguminosae) 253, 254, 389 Ferrisia virgata, 179 Phenacoccus gossypii, 248 Rhizoecus falcifer, 389 Acacia farnesiana (Leguminosae), 181 Acalypha sp. (Euphorbiaceae), 181 Ferrisia virgata, 179 1

Acanthus mollis (Acanthaceae), 253 Phenacoccus gossypii, 248 Acer sp. (Aceraceae), 303 Heliococcus stachyos, 186 Pseudococcus longispinus, 303 Acer platanoides (Aceraceae), 186 Achillea sp. (Compositae) 277, 280 Ferrisia virgata, 179 Phenacoccus solenopsis, 277 Rhizoecus eluminatus, 387 Rhizoecus gracilis, 392 Rhizoecus pritchardi, 400 Spilococcus pressus, 442 Achillea ageratifolia (Compositae), 387, 400 Achillea millefolium (Compositae) 392, 394, 442, 444 Achillea umbellata (Compositae), 181 Achyranthes indica (Amarantaceae), 181 Ferrisia virgata, 179 Acorus sp. (Araceae), 283 Planococcus citri, 280

Some plants do not have a common name. 501

Adenostoma sp. (Rosaceae), 56, 59 Anisococcus adenostomae, 56 Anthelococcus simondsi, 71 Heliococcus adenostomae, 183 Heliococcus atriplicis, 183 Heliococcus stachyos, 186 Adenostoma fasciculatum (Rosaceae), 5, 56, 58, 59, 71, 72, 73, 183, 184, 186, 188 Adiantum sp. (Polypodiaceae), 5, 383, 384, 385, 389, 400, 402 Rhizoecus cyperalis, 383 Rhizoecus falcifer, 389 Rhizoecus pritchardi, 400 African lily. See Agapanthus africanus (Liliaceae) African violet, see Saintpaulia Agapanthus sp. (Liliaceae), 391 Chorizococcus lounsburyi, 103 Rhizoecus falcifer, 389 Agapanthus africanus (Liliaceae), 103

Agapanthus umbellatus (Liliaceae), 103 Agathaea coelestis (^Felicia amelloides) (Compositae), 253 Phenacoccus gossypii, 248 Agathis sp. (Araucariaceae), 206, 208 Nipaecoccus aurilanatus, 206 Agavaceae, 471 Agave sp. (Agavaceae), 111, 112, 113, 465 Chorizococcus reducta, 111 Trionymus diminutus, 465 Agave family. See Agavaceae Aglaonema sp. (Araceae), 316 Pseudococcus obscurus, 313 Agoseris sp. (Compositae), 244, 247 Chorizococcus wilkeyi, 119 Phenacoccus eschscholtziae, 244 Agoseris grandiflora (Compositae), 119, 120, 121

502

HOST

INDEX

Agropyron sp. (Gramineae), 130, 189, 490, 491 Amonostherium lichtensioides, 52 Cryptoripersia salina, 130 Heterococcus arenae, 189 Heterococcus pulverarius, 191 Trionymus winnemucae, 490 Agropyron repens (Gramineae), 191 Agropyron spicatum (Gramineae), 54, 130 Agrostis sp. (Gramineae), 389, 391, 394, 395, 457 Rhizoecus falcifer, 389 Rhizoecus graminis, 394 Tridiscus multiorbis, 457 Trionymus smithii, 485 Agrostis sp. prob, rubra (Gramineae), 485, 487 Ajuga sp. (Labiatae), 391 Rhizoecus falcifer, 389 Alchemilla orbiculata (Rosaceae) 372 Rhizoecus associatus, 372 alder. See Alnus alfalfa. See Medicago sativa alkali heath. See Frankenia grandifolia alkali mallow. See Sida alkali weed. See Cressa eretica Allenrolfea occidentalis (Chenopodiaceae ), 455, 457 Tridiscus distichlii, 455 almond. See Prunus amygdalus Alnus sp. (Betulaceae), 309 Pseudococcus maritimus, 306 Alocasia sp. (Araceae), 181 Ferrisia virgata, 179 Aloe sp. (Liliaceae), 104, 105, 481, 483 Chorizococcus lounsburyi, 103 Trionymus myersi, 481 Alopecurus sp. (Gramineae), 191 Heterococcus pulverarius, 191 Althaea sp. (Malvaceae) 252, 254, 277, 316, 442, 444 Phenacoccus gossypii, 248 Phenacoccus solani, 273 Phenacoccus solenopsis, 277 Pseudococcus obscurus, 313 Spilococcus pressus, 442 alum root. See Heuchera Althaea rosea (Malvaceae), 442, 444 Spilococcus pressus, 442 amaranth. See Amaranthus Amaranthus sp. (Amaranthaceae), 275, 316 Phenacoccus solani, 273 Pseudococcus obscurus, 313

Amaryllis sp. (Amaryllidaceae) 103, 105, 282, 317 Chorizococcus lounsburyi, 103 Planococcus citri, 280 Pseudococcus obcurus, 313 Amazon lily. See Eucharis Ambrosia sp. (Compositae), 231, 234, 252, 253, 254, 273, 275, 277, 279 Phenacoccus defectus, 231 Phenacoccus gossypii, 248 Phenacoccus solani, 273 Phenacoccus solenopsis, 277 Ambrosia artemisifolia (Compositae), 277 Ambrosia psilostachya (Compositae), 275, 276, 277 Amelanchier sp. (Rosaceae), 231 Phenacoccus dearnessi, 231. Amelanchier pallida (Rosaceae), 231, 232 American beach grass. See Ammophila breviligulata Ammophila arenaria (Gramineae) 487, 489 Trionymus utahensis, 487 Ammophila breviligulata (Gramineae), 368 Radicoccus kelloggi, 368 Amelopsis quinquefolia (Vitaceae), 85 Chnaurococcus trifolii, 83 Amsinckia sp. (Boraginaceae) 244, 247 Phenacoccus eschscholtziae, 244 Amygdalus, 231. See also Prunus Ananas comosus (Bromeliaceae) 160, 280, 303 Dysmicoccus brevipes, 160 Planococcus citri, 280 Pseudococcus longispinus, 303 Ananas sativus ( = A . comosus) (Bromeliaceae), 280 Anaphalis margaritacea (Compositae) 262, 264 Phenacoccus lotearum, 262 Anchusa myosotidiflora ( = B r u n n e r a macrophylla) (Boraginaceae), 305 Pseudococcus longispinus, 303 Anemone sp. (Ranunculaceae) 318, 389 Pseudococcus obscurus, 313 Rhizoecus falcifer, 389 Anemone japonica (Ranunculaceae), 391 Annona sp. (Annonaceae), 181, 2 1 0

Dysmicoccus brevipes, 160 Ferrisia virgata, 179 Nipaecoccus nipae, 208 Planococcus citri, 280 Annona muricata (Anonaceae) 160 Annona squamosa (Annonaceae), 280 ant nests, 130, 132, 133, 230, 279, 302 Cryptoripersia salina, 130 Cryptoripersia tubulata, 132 Phenacoccus colemani, 227 Phenacoccus solenopsis, 277 Pseudococcus longisetosus, 300 antelope brush. See Purshia Antennaria argentea (Compositae), 276 Phenacoccus solani, 273 Anthemis sp. (Compositae), 52, 55 Amonostherium lichtensioides, 52 Rhizoecus falcifer, 389 Anthemis tinctoria (Compositae), 391 Anthoxanthum odoratum (Gramineae), 469, 471 Trionymus festucae, 469 Apium sp. (Umbelliferae), 160 Dysmicoccus brevipes, 160 Phenacoccus helianthi, 257 Planococcus citri, 280 Apium graveolens (Umbelliferae), 253, 283 Aplopappus sp. (Compositae), 426 Chorizococcus abortivus, 89 Spilococcus eriogoni, 423 Aplopappus pinifolius (Compositae) 89, 90, 91 apple. See Malus aquatic palm. See Nipa fruticans Aquilegia sp. (Ranunculaceae), 276, 295, 389, 392 Phenacoccus solani, 273 Pseudococcus fragilis, 295 Rhizoecus falcifer, 389 Aralia sp. (Araliaceae), 254, 391 Phenacoccus gossypii, 248 Rhizoecus falcifer, 389 Araucaria sp. (Araucariaceae) 206, 208 Dymicoccus ryani, 170 Nipaecoccus aurilanatus, 206 Araucaria bidwillii (Araucariaceae) 206, 208 Araucaria excelsa, 5, 170, 171, 172, 206, 207, 208 Araucariaceae, 168 arborvitae. See Thuja

Arbutus sp. (Ericaceae) 306, 308 Pseudococcus maritimus, 306 arbute tree. See Arbutus Arctium lappa (Compositae), 227 Phenacoccus colemani, 227 Arctostaphylos sp. (Ericaceae), 8, 186, 188, 329, 331, 332, 333, 335, 336, 337, 363, 365, 400 Anthelococcus simondsi, 71 Heliococcus stachyos, 186 Phenacoccus colemani, 227 Pseudococcus fragilis, 295 Puto albicans, 329 Puto arctostaphyli, 335 Puto yuccae, 363 Rhizoecus pritchardi, 400 Arctostaphylos mariposa (Ericaceae), 332 Arctostaphylos pajaroensis (Ericaceae) 297 Arctostaphylos rudis (Ericaceae), 71, 72, 73, 227, 230 Arctostaphylos viscida (Ericaceae), 337 Areca lutescens (Palmaceae), 162 Dysmicocus brevipes, 160 areca palm. See Areca lutescens Arenaria verna (Caryophyllaceae), 276 Phenacoccus solani, 273 Aristida sp. (Gramineae), 189, 191 Heterococcus arenae, 189 Armeria sp. (Plumbaginaceae), 315, 318 Pseudococcus longisetosus, 300 Pseudococcus obscurus, 313 Armeria vulgaris ( = A . maritima var. purpurea) (Plumbaginaceae), 300, 302 Arrhenatherum elatius (Gramineae), 191 Heterococcus pulverarius, 191 arrow root. See Maranta arrow weed. See Pluchea sericea Artemisia sp. (Compositae), 16, 52, 54, 55, 219, 221, 322, 324 Amonostherium litchtensioides, 52 Anisococcus didymus, 61 Phenacoccus artemisiae, 219 Phenacoccus solani, 273 Pseudococcus obscurus, 313 Pseudococcus sorghiellus, 322 Puto pacificus, 354

HOST Artemisia sp. (Cont.) Rhizoecus californicus, 381 Rhizoecus gracilis, 392 Spilococcus corticosus, 421 Spilococcus eriogoni, 423 Spilococcus geraniae, 427 Spilococcus keiferi, 434 Tridiscus multiorbis, 457 Artemisia absinthium (Compositae) 273, 276, 457, 459 Artemisia californica (Compositae), 54, 55, 59, 62, 63, 219, 220, 221, 276, 423, 426 Artemisia douglasiana (Compositae), 381, 383, 427, 428, 429 Artemisia dracunculus (Compositae) 315, 318 Artemisia frigida (Compositae), 52 Artemisia heterophylla (Compositae), 273, 277, 434, 435, 436 Artemisia pycnocephala (Compositae), 354, 356, 357 Artemisia tridentata (Compositae), 54, 221, 392, 394, 421, 422, 423 Artocarpus altilis (Moraceae), 210 Nipaecoccus nipae, 208 Arundinaria sp. (Gramineae), 286 Antonina pretiosa, 78 Pseudantonina arundinariae, 286 Arundinaria gigantea (Gramineae), 78, 79 Arundo donax (Gramineae), 113, 115, 148, 149, 150 Chorizococcus rostellum, 113 Distichlicoccus arundinis, 148 Asclepiadaceae 95 Asclepias sp. (Asclepiadaceae), 254 Chorizococcus brevicruris, 93 Ferrisia virgata, 179 Phenacoccus gossypii, 248 Asclepias curassavica (Asclepiadaceae) 7, 181 ash. See Fraxinus Asparagus sp. (Liliaceae), 210, 254, 280, 303, 423, 426 Nipaecoccus nipae, 208 Phenacoccus gossypii, 248 Planococcus citri, 280 Pseudococcus longispinus, 303 Spilococcus eriogoni, 423 Asplenium sp. (Polypodiaceae), 315, 317 Pseudococcus obscurus, 313 Aster adscendens (Compositae) 273, 276 Phenacoccus solani, 273

Aster chilensis (Compositae), 442, 444 Spilococcus pressus, 442 Aster spinosus (Compositae), 442, 444 Spilococcus pressus, 442 Astragalus miguelensis (Leguminosae), 363, 365 Puto yuccae, 363 Astrophytum sp. (Cactaceae), 419 Spilococcus cactearum, 416 Atriplex sp. (Chenopodiaceae), 194, 195, 277, 279, 337, 338, 339, 363, 392,436 Anisococcus imperialis, 65 Heliococcus atriplicis, 183 Humococcus atriplicis, 194 Humococcus inornatus, 199 Phenacoccus solenopsis, 277 Puto atriplicis, 337 Puto echinatus, 347 Puto yuccae, 363 Rhizoecus gracilis, 392 Spilococcus atriplicis, 414 Spilococcus larreae, 436 Atriplex canescens (Chenopodiaceae), 65, 66, 67, 194, 416 Atriplex hymenelytra (Chenopodiaceae), 194, 199 Atriplex lentiformis (Chenopodiaceae), 183, 185, 186

Atriplex polycarpa (Chenopodiaceae), 279 Atriplex semibaccata (Chenopodiaceae), 8, 347, 349, 350 Aucuba sp. (Cornaceae), 254, 318 Phenacoccus gossypii, 248 Pseudococcus obscurus, 313 Aucuba japonica (Cornaceae), 253 Audibertia sp. (=Salvia sp.) (Labiatae), 52, 54 Amonostherium lichtensioides, 52 Anisococcus crawii, 59 Anisococcus didymus, 61 Phenacoccus eschscholtziae, 244 Phenacoccus gossypii, 248 Phenacoccus helianthi, 257 Puto yuccae, 363 Audibertia polystachya (=Salvia polystachya) (Labiatae), 59 Australian pink vine, 254 Australian saltbrush, see Atriplex semibaccata Avena barbata (Gramineae), 487 Trionymus utahensis, 487 Avena fatua (Gramineae), 113, 115, 255, 257

Chorizococcus rostellum, 113 Phenacoccus graminosus, 255 avens. See Geum avocado, 7, 12. See also Persea Azalea sp. (Ericaceae), 126, 127 Crisicoccus azaleae, 125 Pseudococcus obscurus, 313 Azalea indica (Ericaceae), 316 Baccharis sp. (Compositae), 81, 107, 109, 231, 234, 242, 244, 257, 259, 260, 279, 416 Cataenococcus olivaceaus, 81 Chorizococcus polyporus, 107 Phenacoccus defectus, 231 Phenacoccus eriogoni, 242 Phenacoccus helianthi, 257 Phenacoccus solenopsis, 277 Pseudococcus obscurus, 313 Puto yuccae, 363 Spilococcus atriplicis, 414 Baccharis pilularis (Compositae) 315, 317, 318 Baccharis viminea (Compositae) 363, 365 bamboo (Gramineae), 74, 75, 76, 78, 80, 286, 287, 477, 478, 479 Antonina crawii, 74 Antonina pretiosa, 78 Pseudantonina arundinariae, 286 Trionymus mocus, 477 Bambusa sp. (Gramineae), 80 Antonina pretiosa, 78 banana. See Musa banyan. See Ficus benghalensis bark, 18, 168, 221, 293, 298, 343, 350, 359, 360, 394 Dysmicoccus quercicolus, 166 Phenacoccus artemisiae, 219 Pseudococcus dispar, 291 Pseudococcus fragilis, 295 Puto cupressi, 343 Puto laticribellum, 350 Puto pricei, 357 Puto profusus, 360 Rhizoecus gracilis, 392 Barley. See Hordeum vulgare Batis sp. (Batidaceae), 315, 317, 347, 348, 363, 365 Pseudococcus obscurus, 313 Puto echinatus, 347 Puto yuccae, 363 bean, 250, 253. See also Phaseolus

INDEX

503

Bebbia juncea var. asper (Compositae) 354, 355 Puto nulliporus, 354 beech family. Sec Fagaceae beefwood, she-oak. See Casuarina bee sage. See Hyptis beet. See Beta Begonia sp. (Begoniaceae), 253, 283, 315, 316, 317 Pseudococcus obscurus, 313 Phenacoccus gossypii, 248 Planococcus citri, 280 beneath paper bands, 461 bent grass. See Agrostis Berberis vulgaris (Berberidaceae), 298 Pseudococcus fragilis, 295 Berlese funnel, 144, 366, 376, 402 Bermuda grass, 7, 8. See also Cynodon dactylon Bertholletia sp. (Lecythidaceae), 257 Cataenococcus olivaceus, 81 Phenacoccus helianthi, 257 Bertholletia sericea (Lecythidaceae), 81 Beta saccharifera (Chenopodiaceae), 276 Phenacoccus solani, 273 Beta vulgaris (Chenopodiaceae), 283 Planococcus citri, 280 Betony. See Stachys bullata bindweed, morning glory. See Convolvulus bird-of-paradise flower. See Strelitzia bird's foot trefoil. See Lotus black sage, 61 Blackburnia umbraculifera (Palmaceae), 389 Rhizoecus kondonis, 394 blue daisy or blue marguerite. See Agathaea coelestis (=Felicia amelloides) bluegrass. See Poa blood flower. See Asclepias curassavica blossom, 61, 63 blueberry, cranberry. See Vaccinium bluegrass, 8, 189. See also Poa douglasii Heterococcus arenae, 189 Heterococcus pulverarius, 191 board (under), 19, 85, 115, 257 Chnaurococcus trifolii, 83 Chorizococcus rostellum, 113 Phenacoccus graminosus, 255 Borage family. See Boraginaceae Boraginaceae, 188 boring beetle tunnels, 115, 116, 117

504

HOST

INDEX

Boston fern, 316. See also Nephrolepis bostoniensis Bouganvillea sp. (Nyctaginaceae), 280 Planococcus citri, 280 Bouteloua sp. (Gramineae), 257, 273 Phenacoccus helianthi, 257 Phenacoccus solani, 273 Bouvardia sp. (Rubiaceae), 254 Phenacoccus gossypii, 248 box. See Buxus boxthorn. See Lycium brambles, including blackberries, dewberries, raspberries. See Rubus branches, 16, 93, 164, 416 Brassavola nodosa (Ochidaceae), 311, 313 Pseudococcus microcirculus, 311 Brassica sp. (Cruciferae), 295 Pseudococcus fragilis, 295 Brayodendron texanum (Ebenaceae), 81 Cataenococcus olivaceus, 81 Brazilnut. See Bertholletia breadfruit. See Artocarpus altilis brome grass. See Bromus Bromeliaceae, 160 Bromus sp. (Gramineae), 113, 115, 144, 147, 457, 487, 489 Chorizococcus rostellum, 113 Cryptoripersia salina, 130 Discococcus spectabilis, 144 Heterococcus pulverarius, 191 Misericoccus arenarius. 204 Phenacoccus graminosus, 255 Rhizoecus browni, 376 Tridiscus multiorbis, 457 Trionymus utahensis, 487 Bromus arenarius (Gramineae), 204 Bromus carinatus (Gramineae), 191 Bromus catharticus (Gramineae), 379 Bromus sp. prob, orcuttianus (Gramineae), 489 Bromus rigidus (Gramineae) 255, 257 Bromus rubens (Gramineae), 144, 146, 147, 204, 205 Bromus sterilis (Gramineae), 191 Bromus tectorum (Gramineae), 130, 132 broom. See Cytisus or Genista broom rape. See Orobanche buck brush. See Ceanothus cuneatus

bugle weed. See Ajuga bugloss. See Anchusa myosotidiflora (=Brunnera macrophylla) bulb canarygrass. See Phalaris tuberosa bulbs, 105, 275, 308, 317 Chorizococcus lounsburyi, 103 Phenacoccus solani, 273 Pseudococcus maritimus, 306 Psedococcus obscurus, 313 bunch grass (Gramineae), 130, 366, 367, 368, 370 bunya-bunya. See Araucaria bidwillii burro-weed. See Franseria dumosa bursage. See Franseria Bursera microphylla (Burseraceae), 257 Phenacoccus helianthi, 257 bush arrowleaf. See Hofmeisteria pleuriseta Buxus sp. (Buxaceae), 298, 389, 391 Pseudococcus fragilis, 295 Rhizoecus falcifer, 389 Buxus africana (Buxaceae), 317 Buxus sempervirens (Buxaceae), 389, 391 cacao, 10. See also Theobroma cacao cactus, 7, 182, 201, 202, 203, 305, 315, 316, 317, 318, 418, 419. See also Cactus and several genera Ferrisia virgata, 179 Hypogeococcus spinosus, 201 Pseudococcus longispinus, 303 Pseudococcus obscurus, 313 Spilococcus cactearum. 416 Cactus sp. (Cactaceae), 115, 276, 277, 279, 280, 398, 399, 418 Chorizococcus rostellum, 113 Phenacoccus solani, 273 Phenacoccus solenopsis, 277 Planococcus citri, 280 Rhizoecus leucosomus, 398 Spilococcus cactearum, 416 calabush tree. See Crescentia Caladium sp. (Araceae), 303 Pseudococcus longispinus, 303 Calendula sp. (Compositae), 252, 254, 275 Phenacoccus gossypii, 248 Phenacoccus solani, 273

California mugwort. See Artemisia heterophylla California nutmeg. See Torreya californica California pepper tree. See Schinus molle California poppy. See Eschscholtzia californica California sagebrush. See Artemisia californica California tea. See Psoralea physodes Calla sp. (Araceae), 298 Pseudococcus fragilis, 295 calla lily, 105, 308, 391. See also Zantedeschia Calluna sp. (Ericaceae), 253 Phenacoccus gossypii, 248 Calonyction sp. (=Ipomoea) (Convolvulaceae), 253 Phenacoccus gossypii, 248 caltrop. See Kallstroemia or Tribulus Camellia sp. (Theaceae), 317 Pseudococcus obscurus, 313 camomile. See Anthemis canary grass, 8, see Phalaris cane. See Arundinaria cane grass. See Arundo donax Canna sp. (Cannaceae) 160, 181, 2 1 0

Dysmicoccus brevipes, 160

Ferrisia virgata, 179 Nipaecoccus nipae, 208 Canon oak. See Quercus chrysolepsis Canthium odoratum (Rubiaceae), 303 Pseudococcus longispinus. 303 cape marigold. See Dimorphotheca Capsicum frutescens longum (Solanaceae), 252 Phenacoccus gossypii, 248 Caralluma nebrowni (Asclepiadaceae), 7, 93, 94, 95 Chorizococcus brevicruris, 93 Carex sp. (Cyperaceae), 137, 138, 306, 389, 394, 398, 400 Discococcus caricis, 137 Pseudococcus maritimus, 306 Rhizoecus falcifer, 389 Rhizoecus kondonis, 394 Rhizoecus leucosomus, 398 Trionymus caricis, 463 Carex integrata (Cyperaceae), 137 Carex tribuloides (Cyperaceae), 463 carob, St. Johns-bread. See Ceratonia siliqua Carya sp. (Juglandaceae), 254

Phenacoccus gossypii, 248 Cassia sp. (Leguminosae), 257 Ferrisia virgata, 179 Phenaoccus helianthi, 257 Cassia guadichaudii (Leguminosae), 181 Castanea sp. (Fagaceae), 297 Pseudococcus fragilis, 295 Castilleia sp. (Schrophulariaceae), 85, 168, 169, 227, 230, 276 Chnaurococcus trifolii, 83 Dysmicoccus racemus, 168 Phenacoccus colemani, 227 Phenacoccus solani, 273 Pseudococcus longisetosus, 300 Castilleia applegateii (Scrophulariaceae), 273 Castilleia foliolosa (Scrophulariaceae), 300, 302 Casuarina ( Casuarinaceae ), 74 Sphaerococcus casuarinae, 74 Catalpa sp. (Bignoniaceae), 186, 306 Heliococcus stachyos, 186 Pseudococcus maritimus, 306 Cattleya sp. (Orchidaceae), 298, 300, 311, 313, 316 Pseudococcus importatus, 298 Pseudococcus microcirculus, 311 Pseudococcus obscurus, 313 Ceanothus sp. (Rhamnaceae), 188, 298, 305, 306, 421 Heliococcus stachyos, 186 Pseudococcus fragilis, 295 Pseudococcus longispinus, 303 Pseudococcus maritimus, 306 Puto albicans, 329 Puto yucca, 363 Rhizoecus bicirculus, 374 Rhizoecus californicus, 381 Spilococcus ceanothi, 419 Spilococcus quercinus, 446 Ceanothus cuneatus (Rhamnaceae), 329, 331, 333, 381, 448 Ceanothus griseus var. horizontalis (Rhamnaceae), 298 Ceanothus oliganthus (Rhamnaceae), 363 Ceanothus prostratus (Rhamnaceae), 419, 420, 421 Ceanothus pumilus (Rhamnaceae), 374, 375, 419, 421 Cedrus sp. (Cupressaceae), 316

HOST Cedrus sp. (Cont.) Pseudococcus longispinus, 303 Pseudococcus obscurus, 313 Cedrus deodora (Cupressaceae), 305, 317 Cedar, 178, 412. See also Cedrus Ehrhorni cupressi, 176 Spilococcus andersoni, 412 celery. See Apium Celosia floribunda (Amaranthaceae), 257, 259, 333 Phenacoccus helianthi, 257 Puto ambiguus, 333 Celtis sp. (Ulmaceae), 225, 306 Phenacoccus celtisifoliae, 225 Pseudococcus maritimus, 306 Rhizoecus kondonis, 394 Celtis occidentalis (Ulmaceae), 397 cement walls, 115 Chorizococcus rostellum, 113 Cenchrus sp. (Gramineae), 113, 115 Chorizococcus rostellum, 113 Centaurea diluta (Compositae), 276 Phenacoccus lotearum, 262 Phenacoccus solani, 273 Centaurea melitensis (Compositae), 262 Centaurea solstitialis (Compositae), 276 Centromadia pungens (Compositae), 279 Phenacoccus solenopsis, 277 cerambycid boring tunnels, 17, 221 Ceratonia siliqua (Leguminosae), 282, 305, 315, 317 Planococcus citri, 280 Pseudococcus longispinus, 303 Pseudococcus obscurus, 313 Cercis canadensis (Leguminosae), 315, 318 Pseudococcus obscurus, 313 Cereus formosus monstrosus (Cactaceae), 418 Spilococcus cactearum, 416 Cereus peruvianus (Cactaceae), 418, 419 Spilococcus cactearum, 416 ceriman. See Monstera Cevallia sinuata (Compositae), 211 Phenacoccus solenopsis, 111

Chaenactis tenuifolia (Compositae), 262 Phenacoccus lotearum, 262 Chaetochloa caudata (Gramineae), 485 Trionymus smithii, 485 Chamaebatia australis (Rosaceae), 365 Puto yuccae, 363 Chamaebatia foliolosa (Rosaceae), 365 Puto yuccae, 363 Chamaecrista procumbens (Leguminosae), 322 Pseudococcus sorghiellus, 322 Chamaecyparis lawsoniana (Cypressaceae), 317 Pseudococcus obscurus, 313 Chamaecyparis lawsoniana var. wisselii (Cupressaceae), 164, 165, 166 Dysmicoccus pinicolus, 164 Chamaecyparis nootkatensis (Cupressaceae), 412, 414 Spilococcus andersoni, 412 chamise. See Adenostoma fasiculatum chaparral, 16, 56, 59, 381, 382, 383, 392, 393, 394 Anisococcus adenostomae, 56 Rhizoecus californicus, 381 Rhizoecus gracilis, 392 Chenopodiaceae, 231, 234, 277, 279, 392, 394 Chenopodium sp. (Chenopodiaceae), 254, 273, 275, 276, 347, 350 Phenacoccus gossypii, 248 Phenacoccus solani, 273 Puto echinatus, 347 chestnut. See Castanea China lily, 105 Chilopsis linearis (Bignoniaceae), 254 Phenacoccus gossypii, 248 Chloris gay ana (Gramineae), 76 Antonina graminis, 76 Chloris inflata (Gramineae), 160 Dysmicoccus brevipes, 160 Choisya sp. (Rutaceae), 253, 283, 298 Phenacoccus gossypii, 248 Planococcus citri, 280 Pseudococcus fragilis, 295 Choisya ternata (Rutaceae), 295, 297, 298 Christmas berry, toyon. See Photinia arbutifolia Chrysanthemum sp. (Compositae), 7, 181, 252, 253, 254, 316, 389, 391, 397 400, 402 Ferrisia virgata, 179 Phenacoccus gossypii, 248

Pseudococcus obscurus, 313 Rhizoecus falcifer, 389 Rhizoecus kondonis, 394 Rhizoecus pritchardi, 400 Chrysanthemum frutescens (Compositae), 389, 391 Chrysothamnus sp. (Compositae), 268, 271, 275 Phenacoccus multisetosus, 268 Phenacoccus solani, 273 Cistus sp. (Cistaceae), 389 Rhizoecus falcifer, 389 citrus, 6, 7, 12. See also Citrus Citrus sp. (Rutaceae), 181, 252, 253, 254, 280, 282, 283, 284, 285, 297, 303, 305, 315, 316, 317, 318, 365, 394, 397 Ferrisia virgata, 179 Phenacoccus gossypii, 248 Phenacoccus helianthi, 257 Planococcus citri, 280 Planococcus kraunhiae, 283 Pseudococcus fragilis, 295 Pseudococcus longispinus, 303 Pseudococcus obscurus, 313 Puto yuccae, 363 Rhizoecus falcifer, 389 Rhizoecus kondonis, 394 Citrus limon (Rutaceae), 281, 282, 283, 305, 316, 317, 363, 366 Citrus nobilis var. unshi (Rutaceae), 283 Citrus paradisi (Rutaceae), 283, 305, 317 Citrus sinensis (Rutaceae), 257, 259, 282, 283, 295, 296, 298, 315, 317, 318, 389 Claytonia lanceolata (Portulacaceae), 379 Rhizoecus browni, 376 Clematis sp. (Ranunculaceae), 316 Pseudococcus obscurus, 313 Clematis armandi (Ranunculaceae), 317 Clivia sp. (Amaryllidaceae), 103, 105 Chorizococcus lounsburyi, 103 clover, 85. See also Trifolium coast dunes, 93 Chorizococcus abroniae, 91 Coast live oak. See Quercus agrifolia Coccoloba sp. (Polygonaceae), 210 Nipaecoccus nipae, 208 cockscomb. See Celosia coconut. See Cocos Cocos nucifera (Palmaceae), 280

INDEX

505

Planococcus citri, 280 Codiaeum sp. (Euphorbiaceae), 257, 280, 389 Phenacoccus helianthi, 257 Planococcus citri, 280 Pseudococcus longispinus, 303 Rhizoecus falcifer, 389 Codiaeum variegatum (Euphorbiaceae), 303 Coffea sp. (Rubiaceae), 160 Rhizoecus falcifer, 389 Coffea liberica (Rubiaceae) 9, 389 coffee. See Coffea coles, mustards. See Brassica coleus. See Coleus Coleus sp. (Labiatae), 5, 6, 252, 276, 280, 282, 283 Phenacoccus gossypii, 248 Phenacoccus solani, 273 Planococcus citri, 280 Colocasia sp. (Araceae), 181 Ferrisia virgata, 179 columbine. See Aquilegia common sagebrush. See Artemisia tridentata common wormwood. See Artemisia absinthium Compositae, 5, 85, 238, 240, 276, 277, 347, 350, 423, 426 Composite family. See Compositae cone flower. See Rudbeckia conifer, 16, 18 Conium maculatum (Umbelliferae), 253, 295, 297 Phenacoccus gossypii, 248 Pseudococcus fragilis, 295 Convolvulus arvensis (Convolvulaceae), 389 Rhizoecus falcifer, 389 copperleaf. See Acalypha Coprosoma sp. (Rubiaceae), 253 Phenacoccus gossypii, 248 Cordyline sp. (Agavaceae), 211 Nipaecoccus nipae, 208 Cordyline australis (Agavaceae), 465 Trionymus diminutus ?, 465 Cordyline terminalis (Agavaceae), 303 Pseudococcus longispinus, 303 coral tree. See Erythrina Coreopsis californica (Compositae), 244, 247 Phenacoccus eschscholtziae, 244 corms, 317 Pseudococcus obscurus, 313 corn, 276. See also Zea Coryphantha andrea (Cactaceae), 419 Spilococcus cactearum, 416

506

HOST

INDEX

Coryphantha viviparia (Cactaceae), 182 Ferrisia virgata, 179 Cotoneaster sp. (Rosaceae), 297, 316 Pseudococcus fragilis, 295 Pseudococcus obscurus, 313 cotton, 7, 252. See also Gossypium Cotyledon sp. ( = D u d l e y a sp.) (Crassulaceae), 333 333 Puto ambiguus, 333 cowpea. See Vigna sinensis crabgrass. See Digitaria Crassula sp. (Crassulaceae), 316, 442, 444 Pseudococcus obscurus, 313 Spilococcus pressus, 442 Crassulaceae, 277 Phenacoccus solani, 273 Crataegus sp. (Rosaceae), 397, 398 Rhizoecus kondonis, 394 creeper. See Parthenocissus creosote bush. See Larrea Crescentia sp. (Bignoniaceae), 181 Ferrisia virgata, 179 Cressa eretica (Convolvulaceae), 423, 427 Spilococcus eriogoni, 423 Crinum sp. (Amaryllidaceae), 103 Chorizococcus lounsburyi, 103 Crotalaria sp. (Leguminosae), 252 Phenacoccus gossypii, 248 Croton sp. (Euphorbiaceae), 181 Ferrisia virgata, 179 Spilococcus eriogoni, 423 Croton californicus (Euphorbiaceae), 426, 427 crown, 5, 16, 17, 19 Cryophytum sp. (Aizoaceae), 315, 317 Pseudococcus obscurus, 313 Cryptantha sp. (Boraginaceae), 247, 407, 408 Phenacoccus eschscholtziae, 244 Scaptococcus californicus, 407 Cryptantha angustifolia (Boraginaceae), 244, 247 Cucumis melo (Cucurbitaceae), 280 Planococcus citri, 280 Cucurbita sp. (Cucurbitaceae), 282, 305 Planococcus citri, 280 Pseudococcus longispinus, 303 Cucurbita pepo, 283 Cupressaceae, 168, 414, 421, 446 Cupressus sp. (Cupressaceae), 170, 172, 178,

253, 283, 291, 293, 294, 295, 305, 306, 308, 392, 414 431,434, 450 Dysmicoccus ryani, 170 Ehrhorni cupressi, 176 Phenacoccus gossypii, 248 Planococcus citri, 280 Pseudococcus aberrans, 289 Pseudococcus diversus, 293 Pseudococcus longispinus, 303 Pseudococcus maritimus, 306 Rhizoecus falcifer, 389 Spilococcus andersoni, 412 Spilococcus implicatus, 431 Spilococcus sequoiae, 448 Cupressus arizonicus (Cupressaceae), 172, 178 Cupressus goveniana (Cupressaceae), 320, 321, 322, 412 Cupressus guadelupensis (Cupressaceae), 172, 178 Cupressus macnabiana (Cupressaceae), 412, 414 Cupressus macrocarpa (Cupressaceae), 170, 171, 172, 176, 177, 343, 344, 412, 414, 431, 433, 434, 450 Cupressus torulosa (Cupressaceae), 170 custard apple. See Annona Cycas sp. (Cycadaceae), 303, 305 Pseudococcus longispinus, 303 Cycas revoluta (Cycadaceae) 303, 304, 305 Cyclamen sp. (Primulaceae), 305 Pseudococcus longispinus, 303 Cydonia sp. (Rosaceae), 315, 318, 461 Pseudococcus obscurus, 313 Trionymus americanus, 461 Cydonia oblonga (Rosaceae), 317 Cynodon dactylon (Gramineae), 76, 78, 113, 114, 115, 262, 389, 392, 455, 457, 467, 469, 487, 488, 489 Antonina graminis, 76 Chorizococcus rostellum, 113 Phenacoccus loterarum, 262 Rhizoecus falcifer, 389 Tridiscus distichlii, 455 Trionymus dolus, 467 Trionymus utahensis, 487 Cyperus sp. (Cyperaceae), 113, 115, 276, 282, 379, 389

Chorizoccus rostellum, 113 Dysmicoccus brevipes, 160 Phenacoccus solani, 273 Planococcus citri, 280 Rhizoecus cacticans, 379 Rhizoecus cyperalis, 383 Rhizoecus falcifer, 389 Cyperus esculentus (Cyperaceae), 113, 115,273, 277 Cyperus rotundus (Cyperaceae), 160, 276, 379, 380, 381 Cyperus tenerrimus (Cyperaceae), 383 cypress, 293. See also Cupressus cypress family, see Cupressaceae Cytisus sp. (Leguminosae), 56, 59 Anisococcus adenostomae, 56 Phenacoccus gossypii, 248 Cytisus hillieri (Leguminosae), 254 Dahlia sp. (Compositae), 316 Pseudococcus obscurus, 313 dalea. See Parosela Dammar pine. See Agathis dandelion. See Taraxacum Danthonia californica (Gramineae), 5, 130 Cryptoripersia salina, 130 Daphne sp. (Thymelaeaceae), 297, 305 Pseudococcus fragilis, 295 Pseudococcus longispinus, 303 date. See Phoenix date palm. See Phoenix dactylifera Datura sp. (Solanaceae), 254 Phenacoccus gossypii, 248 decaying log, 404 Rhizoecus spinosus, 404 deerweed. See Lotus scoparius Delphinium sp. (Ranunculceae), 389 Rhizoecus falcifer, 389 Denbrobium sp. (Orchidaceae), 298, 299, 300, 311, 313 Pseudococcus importatus, 298 Pseudococcus microcirculus, 311 Deschampsia ceaspitosa (Gramineae), 469, 471 Trionymus festucae, 469 desert, 16 desert eriogonum. See Eriogonum deserticola desert thorn. See Lycium desert willow. See Chilopsis linearis Dianthus sp. (Caryophyllaceae), 315, 317, 318

Pseudococcus obscurus, 313 Dicoria canescens (Compositae), 235, 237, 238 Phenacoccus dicoriae, 235 Dieffenbachia sp. (Araceae), 282, 315, 316 Planococcus citri, 280 Pseudococcus obscurus, 313 Digitalis sp. (Scrophulariaceae), 295 Pseudococcus fragilis, 295 Digitaria sp. (Gramineae), 113 Chorizococcus rostellum, 113 Pseudococcus longispinus, 303 Digitaria sanguinalis (Gramineae), 113, 115 303 Dimorphotheca sp. (Compositae), 254 Phenacoccus gossypii, 248 Dioscorea sp. (Dioscoreaceae), 282 Planococcus citri, 280 Diospyros sp. (Ebenaceae), 283, 285, 317 Planococcus kraunhiae, 283 Pseudococcus longispinus, 303 Pseudococcus obscurus, 313 Diospyros virginiana (Ebenaceae), 303 Diplacus sp. (Scrophulariaceae), 253, 345, 346, 365 Heliococcus stachyos, 186 Nipaecoccus aurilanatus, 206 Phenacoccus gossypii, 248 Puto decorosus, 345 Puto yuccae, 363 Diplacus aurantiacus (Scrophulariaceae), 186, 365 Diplacus longiflorus (Scrophulariaceae), 206, 208 Dipsacus sp. (Dipsaceae), 381, 383 Rhizoecus californicus, 381 Distichlis sp. (Gramineae), 130, 131, 154, 213, 253, 453, 454 Cryptoripersia salina, 130 Distichlicoccus salinus, 154 Dysmicoccus timberlakei, 172 Humococcus caritus, 194 Paludicoccus distichlium, 211 Phenacoccus gossypii, 248 Rhizoecus cacticans, 379 Stemmatomerinx decorata, 453 Tridiscus distichlii, 455 Distichlis spicata (Gramineae), 154, 155, 156,

HOST Distichlis spicata (Cont.) 172, 173, 174, 194, 196, 197, 211, 213, 379, 381, 455, 456, 457 Dizygotheca elegantissima (Araliaceae), 315, 317 Pseudococcus obscurus, 313 dock, sorrel. See Rumex Dodecatheon hendersonii (Primulaceae), 379 Rhizoecus browni, 376 Dombeya sp. (Sterculiaceae), 252 Phenacoccus gossypii, 248 Douglas fir. See Pseudotsuga taxifolia Dracaena sp. (Agavaceae), 210, 305, 389 Nipaecoccus nipae, 208 Pseudococcus longispinus, 303 Rhizoecus falcifer, 389 Dracena indivisa (Agavaceae), 305 dracena, see Dracaena Dudleya sp. (Crassulaceae), 333 Puto ambiguus, 333 Rhizoecus leucosomus, 398 Dudleya farinosa (Crassulaceae), 398 Echeveria sp. (Crassulaceae), 231, 234, 238, 239, 240, 244, 273, 276, 316, 379, 381 Phenacoccus defectus, 231 Phenacoccus echeveriae, 238 Phenacoccus eschscholtziae, 244 Phenacoccus solani, 273 Pseudococcus obscurus, 313 Rhizoecus cacticans, 379 Spilococcus cactearum, 416 Echeveria longissima (Crassulaceae), 231, 234, 418 Echeveria rubella (Crassulaceae), 234, 275 Echeveria scheeri (Crassulaceae), 234 Echinocereus sp. (Cactaceae), 201, 203 Hypogeococcus spinosus, 201 Echinomastus johnsonii (Cactaceae), 5, 418 Spilococcus cactearum, 416 Echinopsis sp. (Cactaceae), 315, 316, 418 Pseudococcus obscurus, 313 Spilococcus cactearum, 416 Echinopsus multiplex (Cactaceae), 418 Echinopsus silvestrii (Cactaceae), 419

Echium sp. (Boraginaceae) 254 Phenacoccus gossypii, 248 elder. See Sambucus elephants ear. See Colocasia elephant tree. See Bursera microphylla Eleusine indica (Gramineae), 113 Chorizococcus rostellum, 113 elm. See Ulmus Elsholtzia stauntonii (Labiatae), 254 Phenacoccus gossypii, 248 Elymus sp. (Gramineae), 130, 132, 189, 329, 330, 467, 469, 471, 472, 475, 479, 481?, 481, 485, 487 Cryptoripersia salina, 130 Heterococcus arenae, 189 Puto acirculus, 329 Trionymus dolus, 467 Trionymus frontalis, 471 Trionymus haancheni, 475 Trionymus modocensis, 479 Trionymus smithii, 485 Elymus condensatus (Gramineae) 5, 329, 467, 469, 479, 4 8 1 , 4 8 5 , 4 8 6 , 487 Elymus glaucus (Gramineae), 191, 193, 461, 465, 467?, 469 Elymus triticoides (Gramineae), 191, 463, 464, 465, 473, 474, 475, 480, 481, 485, 487 Elymus virginicus (Gramineae), 461 Encelia sp. (Compositae), 240, 242 Heliococcus stachyos, 186 Phenacoccus colemani, 227 Phenacoccus eremicus, 240 Phenacoccus gossypii, 248 Phenacoccus solenopsis, 277 Puto mimicus, 352 Encelia farinosa (Compositae), 5, 186, 227, 230, 240, 242, 254, 277, 278, 279 Encelia frutescens (Compositae), 279, 352, 353, 354 Enceliopsis sp. (Compositae), 277, 279 Phenacoccus solenopsis, 277 Englemann spruce. See Picea engelmanii English ivy. See Hederá helix Ephedra sp. (Gnetaceae), 63, 65 Anisococcus ephedrae, 63 Ephedra californica (Gnetaceae), 63, 64

Epidendrum sp. (Orchidaceae), 298, 300, 303 Pseudococcus importatus, 298 Epiphylum sp. (Cactaceae), 379, 381 Rhizoecus cacticans, 379 Epithelantha micromeris (Cactaceae), 315, 316 Pseudococcus obscurus, 313 Eremocarpus setigerus (Euphorbiaceae), 442, 444 Spilococcus pressus, 442 Eriastrum densifolium (Polemoniaceae), 339, 342, 365 Puto californicus, 339 Puto yuccae, 363 Erica sp. (Ericaceae), 253, 298 Phenacoccus gossypii, 248 Pseudococcus fragilis, 295 Ericomeria ericoides (Compositae), 298 Pseudococcus fragilis, 295 Ericameria fasciculata (Compositae), 487, 489 Trionymus utahensis, 497 Ericameria pinifolia (Compositae), 89, 90, 91 Chorizococcus abortivus, 89 Erigeron sp. (Compositae), 252, 273, 276, 322, 324 Phenacoccus gossypii, 248 Phenacoccus solani, 273 Pseudococcus sorghiellus, 322 Eriobotrya japónica (Rosaceae), 303, 397 Pseudococcus longispinus, 303 Rhizoecus kondonis, 394 Eriodictyon sp. (Hydrophyllaceae), 423 Puto yuccae, 363 Spilococcus eriogoni, 423 Eriodictyon californicum (Hydrophyllaceae), 365, 423 Eriodictyon tomentosum (Hydrophyllaceae), 423, 426 Eriogonum sp. (Polygonaceae), 67, 96, 97, 98, 99, 107, 109, 186, 188, 223, 224, 227, 230, 242, 244, 247, 259, 262, 271, 272, 273, 385, 386, 387, 416, 423, 424, 426, 427 Anisococcus oregonensis, 67 Chorizococcus coxindex, 96 Chorizococcus polyporus, 107 Chorizococcus senarius, 115 Heliococcus stachyos, 186 Phenacoccus cajonensis, 221 Phenacoccus colemani, 227

INDEX

507

Phenacoccus defectus, 231 Phenacoccus destitutus, 235 Phenacoccus eriogoni, 242 Phenacoccus eschscholtziae, 244 Phenacoccus helianthi, 257 Phenacoccus lotearum, 262 Phenacoccus pauperatus, 271 Pseudococcus maritimus, 306 Puto yuccae, 363 Rhizoecus browni, 376 Rhizoecus disjunctus, 385 Scaptococcus milleri, 407 Spilococcus atriplicis, 414 Spilococcus eriogoni, 423 Eriogonum deflexum brachypodium (Polygonaceae), 242, 244, 247 Eriogonum deserticola (Polygonaceae) 5, 19, 407, 409, 410, 426 Eriogonum fasciculatum (Polygonaceae), 115, 116, 117, 224, 365, 423, 427 Eriogonum fasciculatum var. foliolosum (Polygonaceae), 107, 109, 225 Eriogonum fasciculatum ssp. polifolium (Polygonaceae), 107, 221, 222, 376 Eriogonum inflatum (Polygonaceae), 186, 188, 416, 426 Eriogonum latifolíum (Polygonaceae), 5, 306, 307, 308, 309, 427 Eriogonum latifolium var. saxícola (Polygonaceae), 235, 236, 242, 244, 426 Eriogonum nudum (Polygonaceae), 242, 243, 244, 427?, 427 Eriogonum umbellatum ssp. polyanthum (Polygonaceae), 5, 68, 186, 188, 231, 234 Eriophyllum sp. (Compositae), 277, 279 Chorizococcus wilkeyi, 119 Phenacoccus defectus, 231 Phenacoccus gossypii, 248 Phenacoccus pauperatus, 271 Phenacoccus solenopsis, 277 Puto yuccae, 363 Eriophyllum confertiflorum (Compositae), 231, 233, 234, 253, 271, 273, 363, 365, 366 Eriophyllum lanatum var, grandiflorum (Compositae), 119, 121

508

HOST

INDEX

Erodium sp. (Geraniaceae), 389 Phenacoccus eschscholtziae, 244 Rhizoecus falcifer, 389 Erodium cicutarium (Geraniaceae), 244, 247 Erythea edulis (Palmaceae), 297, 315, 317 Pseudococcus fragilis, 295 Pseudococcus obscurus, 313 Erythrina sp. (Leguminosae), 252 Phenacoccus gossypii, 248 Escallonia rubra (Saxifragaceae), 391 Rhizoecus falcifer, 389 Eschscholtzia sp. (Papaveraceae), 244, 245, 247, 253 Phenacoccus eschscholtziae, 244 Phenacoccus gossypii, 248 Pseudococcus obscurus, 313 Eschscholtzia californica (Papaveraceae), 316 Escobaría tuberculosa (Cactaceae), 398 Rhizoecus leucosomus, 398 Eucharis amazónica (Amaryllidaceae), 275, 276 Phenacoccus solani, 273 Eugenia sp. (Myrtaceae), 295 Pseudococcus fragilis, 295 Euonymus sp. (Celastraceae), 253, 317, 318 Phenacoccus gossypii, 248 Pseudococcus obscurus, 313 Euphorbia sp. (Euphorbiaceae), 160, 231, 234, 275, 277, 280, 282 Dysmicoccus brevipes, 160 Ferrisia virgata, 179 Phenacoccus defectus, 231 Phenacoccus solani, 273 Phenacoccus solenopsis, 277 Planococcus citri, 280 Pseudococcus obscurus, 313 Euphorbia maculata (Euphorbiaceae), 275, 276 Euphorbia pulcherrima (Euphorbiaceae), 181, 282, 306, 315 European beach grass. See Ammophila arenaria European or Common barberry. See Berberís vulgaris evening primrose. See Oenothera evening primrose family. See Onagraceae Fagaceae, 67 false aralia. See Dizygotheca

false cypress. See Chamaecyparis Fatsia sp. (Araliaceae), 254, 317 Phenacoccus gossypii, 248 Pseudococcus obscurus, 313 fern, 16, 19 Ferocactus coloratus (Cactaceae), 182 Ferrisia virgata, 179 fescue. Sei Festuca Festuca sp. (Gramineae), 150. 151. 152, 191, 255, 257 Distichlicoccus californicus, 150 Heterococcus pulverarius, 191 Phenacoccus graminosus, 255 Radicoccus kelloggi, 368 Trionymus festucae, 469 Festuca ovina (Gramineae), 370 Festuca rubra (Gramineae), 191 Festuca scabrella (Gramineae, 469, 470, 471 Ficus sp. (Moraceae), 81, 282, 295, 298, 397 Cataenococcus olivaceus, 81 Ferrisia virgata, 179 Nipaecoccus nipae, 208 Planococcus citri, 280 Pseudococcus fragilis, 295 Pseudococcus longispinus, 303 Rhizoecus kondonis, 394 Ficus benghalensis (Moraceae), 210, 303 Ficus carica (Moraceae), 283 Ficus decora (Moraceae), 305 Ficus palmeri (Moraceae), 181 Ficus pandoratus (Moraceae), 283 fiddleneck. See Amsinckia fig. See Ficus fig marigold. See Mesembryanthemum fir. See Abies firethorn. See Pyracantha flat top. See Eriogonum fasciculatum fleabane. See Erigeron flowering maple. See Abutilon foliage, 16, 17, 18 fountain palm. See Livistona Fouquieria splendens (Fouquieriaceae), 260, 452 Phenacoccus infernalis, 260 Spilococcus townsendi, 452 foxglove. See Digitalis foxtail. See Alopecurus Fragaria sp. (Rosaceae), 282, 389, 391, 397 Planococcus citri, 280

Rhizoecus falcifer, 389 Rhizoecus kondonis, 394 Franciscea sp. ( = B r u n f e l sia) (Solanaceae), 282 Planococcus citri, 280 Frankenia grandifolia (Frankeniaceae), 279 Phenacoccus solenopsis, 277 Franseria sp. (Compositae), 199, 200, 253, 275. 276, 277, 280 Chorizococcus abroniae, 91 Humococcus inornatus, 199 Phenacoccus alleni, 216 Phenacoccus dicoriae, 235 Phenacoccus eremicus, 240 Phenacoccus gossypii, 248 Phenacoccus helianthi, 257 Phenacoccus solani, 273 Phenacoccus solenopsis, 277 Spilococcus keiferi, 434 Spilococcus parvicirculus, 440 Spilococcus ventralis, 450 Franseria acanthicarpa (Compositae), 440, 441, 442 Franseria bipinnatifida (Compositae), 280 Franseria chamissonis (Compositae), 91, 93, 434, 436 Franseria dumosa (Compositae), 5, 199, 216, 219, 235, 238, 240, 241, 242, 248, 249, 257, 259, 450, 451 Fraxinus sp. (Oleaceae), 461, 465?, 467? Trionymus americanus, 461 Trionymus diminutus, 465 fringed sagebrush, see Artemisia frigida Fuchsia sp. (Onagraceae), 254 Phenacoccus gossypii, 248 furze, gorse. See Ulex galingale. See Cyperus Gardenia sp. (Rubiaceae), 253, 282, 283, 316, 391 Phenacoccus gossypii, 248 Planococcus citri, 280 Pseudococcus obscurus, 313 Rhizoecus falcifer, 389 Gardenia intermedia (Rubiaceae), 389 gardens, 9, 10, 14, 16 Garrya sp. (Garryaceae), 227, 229 Phenacoccus colemani, 227 Puto yuccae, 363 Garrya elliptica (Garryaceae), 363 Garrya veatchii (Garryaceae), 363, 365

Gasteria sp. (Liliaceae), 103, 105 Chorizococcus lounsburyi, 103 Gasteria lingua (Liliaceae), 5 Gaultheria shallon (Ericaceae), 357 Puto pacificus, 354 Genista sp. (Leguminosae), 306, 308, 316, 318 Pseudococcus maritimus, 306 Pseudococcus obscurus, 313 geranium (of florists). See Geranium or Pelargonium Geranium sp. (Geraniaceae), 252, 253, 254, 315, 316 Phenacoccus gossypii, 248 Pseudococcus obscurus, 313 Spilococcus geraniae, 427 Geranium robertianum (Geraniaceae), 427, 428 Gerbera sp. (Compositae), 181 Ferrisia virgata, 179 Geum borisi (Rosaceae), 400 Rhizoecus pritchardi, 400 giant bamboo, 76, 78, 79, 80 Antonina crawii, 74 Antonina pretiosa, 78 giant reed. See Arundo donax giant rye grass. See Elymus condensatus Gladiolus sp. (Iridaceae), 315, 317 Pseudococcus obscurus, 313 glasswort. See Salicornia Gleditsia triacanthos (Leguminosae), 306 Pseudococcus maritimus, 306 Gliricida sp. (Leguminosae), 160 Dysmicoccus brevipes, 160 globe mallow. See Sphaeralcea gloxinia, 388. See also Gloxinia sp. (=Sinningia sp.) Gloxinia sp. (Gesneriaceae), 283, 317, 387 Planococcus citri, 280 Pseudococcus obscurus, 313 Rhizoecus eluminatus, 387 goatnut. See Simmondsia chinensis goats beard. See Tragopogon gold dust. See Aucuba japonica goldenrod. See Solidago goldenweed. See Aplopappus golf green, 8

HOST gooseberry, currant. See Ribes goosefoot, pigweed. See Chenopodium goose grass. See Eleusine indica Gossypium sp. (Malvaceae), 160, 181, 252, 277, 426, 442, 444 Dysmicoccus brevipes, 160 Ferrisia virgata, 179 Phenacoccus gossypii, 248 Phenacoccus solenopsis, 277 Spilococcus eriogoni, 423 Spilococcus pressus, 442 Gowen cypress. See Cupressus goveniana Gramineae, 5, 18 grape. See Vitis grapefruit. See Citrus paradisi grapes, 6, 7, 12 grass (Gramineae), 5, 7, 8, 14, 16, 17, 18, 19 great burdock. See Arctium lappa greenhouses 6, 7, 10, 12, 13, 16 Grevillea sp. (Proteaceae), 295 Pseudococcus fragilis, 295 Grindelia robusta (Compositae), 85 Chnaurococcus trifolii, 83 groundsel. See Senecio guava. See Psidium gum plant. See Grindelia Gutierrezia sp. (Compositae), 257, 430, 431 Chorizococcus abortivus, 89 Phenacoccus gossypii, 248 Phenacoccus helianthi, 257 Spilococcus eriogoni, 423 Spilococcus gutierreziae, 429 Spilococcus parkeri, 438 Gutierrezia californica (Compositae), 426, 438, 439, 440? Gutierrezia microcephala (Compositae), 254, 438 Gutierrezia sarothrae (Compositae), 89, 429, 431 Haanchen barley. See Hordeum vulgare hackberry. See Celtis hair grass. See Deschampsia Hamatocactus sp. (Cactaceae), 5 Rhizoecus cacticans, 379 Spilococcus cactearum, 416 Hamatocactus setispinus (Cactaceae), 5, 379, 381, 418

Haplopappus sp. (Compositae), 253, 257, 259, 416, 429 Anisococcus crawii, 59 Humococcus inornatus, 199 Phenacoccus gossypii, 248 Phenacoccus helianthi, 257 Pseudococcus maritimus, 306 Spilococcus atriplicis, 414 Spilococcus gutierreziae, 429 Haplopappus cooperi? (Compositae), 431 Haplopappus ericoides (Compositae), 306, 308 Haplopappus pinifolius (Compositae), 61, 199 Haplopappus venutus vernonioides (Compositae), 254 Haworthia sp. (Liliaceae), 103, 104, 105, 381, 481, 482, 483 Chorizococcus lounsburyi, 103 Rhizoecus cacticans, 379 Spilococcus cactearum, 416 Trionymus myersi, 481 Haworthia margaritifera (Liliaceae), 419 hawthorn. See Crataegus heath. See Erica heather. See Calluna Hebe sp. (Scrophulariaceae), 305, 318 Pseudococcus longispinus, 303 Pseudococcus obscurus, 313 Hedera helix (Araliaceae), 170, 252, 253, 277, 295, 298, 303, 318 Dysmicoccus ryani, 170 Phenacoccus gossypii, 248 Phenacoccus solani, 273 Pseudococcus fragilis, 295 Pseudococcus longispinus, 303 Pseudococcus obscurus, 313 hedge nettle. See Stachys Helenium sp. (Compositae), 276 Phenacoccus solani, 273 Helianthemum scoparium andersoni (Cistaceae), 244, 247 Phenacoccus eschscholtziae, 244 Helianthus sp. (Compositae), 231, 234, 254, 257, 275, 276, 277, 295 Chnaurococcus trifolii, 83 Phenacoccus defectus, 231 Phenacoccus gossypii, 248 Phenacoccus helianthi, 257 Phenacoccus solani, 273

Phenacoccus solenopsis, 277 Pseudococcus fragilis, 295 Helianthus annuus (Compositae), 276 Helianthus niveus (Compositae), 251, 253, 254 Helianthus techrodes (Compositae), 251, 253 Helianthus tuberosus (Compositae), 85, 254 Heliconia sp. (Musaceae), 252 Phenacoccus gossypii, 248 heliotrope. See Heliotropium Heliotropium arborescens (Boraginaceae), 295 Pseudococcus fragilis, 295 Hemizonia rudis (Compositae), 273, 276 Phenacoccus solani, 273 Hesper palm. See Erythea Heteromeles sp. (=Photinia sp.) (Rosaceae), 225 Phenacoccus celtisifoliae, 225 Heterotheca sp. (Compositae), 275, 426 Phenacoccus solani, 273 Spilococcus eriogoni, 423 Heterotheca grandiflora (Compositae), 276 Heuchera sp. (Saxifragaceae), 315, 318, 391 Pseudococcus obscurus, 313 Rhizoecus falcifer, 389 Hibiscus sp. (Malvaceae), 253, 254, 282, 303, 391 Phenacoccus gossypii, 248 Phenacoccus solani, 273 Planococcus citri, 280 Pseudococcus longispinus, 303 Rhizoecus falcifer, 389 Hibiscus esculentus Malvaceae), 275 Hibiscus syriacus (Malvaceae), 253 hickory. See Carya Hilaria sp. (Gramineae), 197 Humococcus hilariae, 197 Hilaria cenchroides (Gramineae), 197 Hilaria rigida (Gramineae), 198, 199 Hippeastrum sp. (Amaryllidaceae, 103, 276 Chorizococcus lounsburyi, 103 Phenacoccus solani, 273 Hofmeisteria pluriseta (Compositae), 266 Phenacoccus megaulus, 266 Holcus lanatus (Gramineae), 191, 379 Heterococcus pulverarius, 191 Rhizoecus cacticans, 379 holly. See Ilex hollyhock. See Althaea rosea

INDEX

509

Homalocephala texensis (Cactaceae), 417, 418 419 Spilococcus cactearum, 416 home, 8 honey locust. See Gleditsia Hordeum sp. (Gramineae), 204, 255, 257, 457, 459 HeterocOccus pulverarius, 191 Misericoccus arenarius, 204 Phenacoccus graminosus, 255 Tridiscus multiorbis, 457 Trionymus haancheni, 475 Hordeum leporinum (Gramineae), 204, 457, 459 Hordeum murinum (Gramineae), 191 Hordeum vulgare (Gramineae), 8, 475, 476, 477 horehound. See Marrubium Horkelia sp. (Rosaceae), 365 Puto yuccae, 363 horsebrush. See Tetradymia horse crippler cactus. See Homalocephala texensis household pests, 8. houseleek, hen-and-chickens. See Sempervivum Howea sp. (Palmaceae), 389 Rhizoecus falcifer, 389 Howea belmoreana (Palmaceae), 389 Hoya sp. (Asclepiadaceae), 283 Planococcus citri, 280 Huernia sp. (Asclepiadaceae), 7, 93, 95 Chorizococcus brevicruris, 93 Huernia macropodia (Asclepiadaceae), 93, 95 Huernia pillansi (Asclepiadaceae), 93, 95 Hugelia densifolium (Polemoniaceae), 339 Puto californicus, 339 Hymenoclea salsola (Compositae), 216, 217, 218, 219, 240, 242, 257, 258, 259, 260 Phenacoccus alleni, 216 Phenacoccus eremicus, 240 Phenacoccus helianthi, 257 Hymenopappus sp. (Compositae), 181 Ferrisia virgata, 179 Hyptis emoryi (Labiatae), 253 Phenacoccus gossypii, 248 Ilex sp. (Aquifoliaceae), 283, 298, 318 Planococcus kraunhiae, 283

510

HOST

INDEX

Ilex sp (Cont.) Pseudococcus fragilis, 295 Pseudococcus obscurus, 313 Impatiens sp. (Balsaminaceae), 282 Planococcus citri, 280 inanimate objects, 19 incense cedar. See Libocedrus decurrens incienso. See Encelia farinosa Indian corn. See Zea mays Indian lettuce. See Montia ink weed, 277, 279 Phenacoccus solenopsis, 277 insectary, 254, 308, 309 Phenacoccus gossypii, 248 Pseudococcus maritimus, 306 iodine bush. See Allenrolfea occidentalis Ipomoea batatas (Convolvulaceae), 282, 283, 315 Planococcus citri, 280 Pseudococcus obscurus, 313 Iris sp. (Iridaceae), 103, 390, 391 Chorizococcus lounsburyi, 103 Rhizoecus falcifer, 389 Irish broom, 318 Pseudococcus obscurus, 313 Ixia sp. (Iridaceae), 253 Phenacoccus gossypii, 248 Jacaranda sp. (Bignoniaceae) 253, 315, 316 Phenacoccus gossypii, 248 Pseudococcus obscurus, 313 Jacobinia pauciflora (Acanthaceae), 253 Phenacoccus gossypii, 248 jade plant, 316 Pseudococcus obscurus, 313 jajoba. See Simmondsia chinensis Japanese wisteria. See Wisteria floribunda Jasminum sp. (Oleaceae), 254, 303, 389, 391, 392 Phenacoccus gossypii, 248 Pseudococcus longispinus, 303 Rhizoecus falcifer, 389 Johnson grass. See Sorghum halepense Joshua tree. See Yucca brevifolia Juglans sp. (Juglandaceae), 291, 293, 295, 306, 308 Pseudococcus dispar, 291 Pseudococcus fragilis, 295 Pseudococcus maritimus, 306 juniper. See Juniperus

Juniperus sp. (Cupressaceae), 107, 108, 109, 121, 122, 123, 144, 147, 170, 172, 178, 291, 324, 366, 376, 379, 412, 431, 434 Chorizococcus polyporus, 107 Chorizococcus wilsoni, 121 Discococcus spectabilis, 144 Dysmicoccus brevipes, 160 Dysmicoccus ryani, 170 Ehrhorni cupressi, 176 Pseudococcus aberrans, 289 Pseudococcus diversus, 293 Pseudococcus sparsus, 324 Pygmaeococcus morrisoni, 366 Rhizoecus browni, 376 Spilococcus andersoni, 412 Spilococcus implicatus, 431 Juniperus californicus (Cupressaceae), 178 Juniperus chinensis (Cupressaceae), 172 Juniperus hibernica (Cupressaceae), 434 Juniperus macnabiana (Cupressaceae), 325, 326 Juniperus sabina fastigiata stricta (Cupressaceae), 172 Juniperus squamata var. meyeri (Cupressaceae), 293, 295 Juniperus torulosa (Cupressaceae), 162, 289, 290, 291 kafir lily. See Clivia Kalanchoe sp. (Crassulaceae), 276 Phenacoccus solani, 273 Kallstroemia sp. (Zygophyllaceae), 259 Phenacoccus helianthi, 257 Phenacoccus solenopsis, 277 Kallstroemia brachystylos (Zygophyllaceae), 277 Kentia sp. (Palmaceae), 5, 209, 210, 211, 298, 389, 391 "Nipaecoccus nipae, 208 Pseudococcus fragilis, 295 Rhizoecus falcifer, 389 Kleiniasp. (Compositae), 379, 381 Rhizoecus cacticans, 379 Rhizoecus falcifer, 389 Kleinia péndula (Compositae), 389 Kniphofia uvaria (Liliaceae), 316 Pseudococcus obscurus, 313

knotweed. See Polygonum Kraunhia floribunda ( = Wisteria floribunda) (Leguminosae), 283 Planococcus kraunhiae, 283 Lachnanthes tinctoria (Haemodoraceae), 322 Pseudococcus sorghiellus, 322 Lantana sp. (Verbenaceae), 5, 7, 113, 115, 180, 181, 252, 253, 254 Chorizococcus rostellum, 113 Ferrisia virgata, 179 Phenacoccus colemani, 227 Phenacoccus gossypii, 248 Phenacoccus solani, 273 Lantana camara (Verbenaceae), 227, 275 larkspur. See Delphinium Larrea sp. (Zygophy 1laceae), 259, 260,438 Phenacoccus helianthi, 257 Spilococcus larreae, 436 Larrea divaricata (=Larrea tridentata) (Zygophyllaceae), 436, 437, 438 Larrea tridentata (Zygophy 1laceae), 436, 437, 438 Lathy rus littoralis (Leguminosae), 91, 93, 426 Chorizococcus abroniae, 91 Spilococcus eriogoni, 423 laurel, sweet bay. See Laurus Laurus nobilis (Lauraceae), 298 Pseudococcus fragilis, 295 Lavandula sp. (Labiatae), 181

Ferrisia virgata, 179 lavender. See Lavandula lawn, 8 Layia glandulosa (Compositae), 244, 247 Phenacoccus eschscholtziae, 244 leaf mold, 9 leather root. See Psoralea macrostachya leaves, 16 Lecythis sp. (Lecythidaceae), 181 Ferrisia virgata, 179 Lepidospartum squamatum (Compositae), 363, 365 Puto yuccae, 363 lemon. See Citrus limon Libocedrus decurrens (Cupressaceae), 178, 350, 352,412,413,414 Ehrhorni cupressi, 176 Puto laticribellum, 350 Spilococcus andersoni, 412 Ligustrum sp. (Oleaceae), 295, 389, 397 Pseudococcus fragilis, 295 Rhizoecus falcifer, 389 Rhizoecus kondonis, 394

lilac. See Syringa Liliaceae, 275 Lilium sp. (Liliaceae), 276 Phenacoccus solani, 273 Lilium longiflorum var. eximium (Liliaceae), 105 Chorizococcus lounsburyi, 103 lily. See Lilium lily family. See Liliaceae lily turf. See Ophiopogon linden. See Tilia litchi. See Nephelium Livistona sp. (Palmaceae), 211 Nipaecoccus nipae, 208 Lobivia sp. (Cactaceae), 379 Rhizoecus cacticans, 379 Lobivia shaferi (Cactaceae), 379 lodgepole pine. See Pinus contorta var. latifolia Lolium sp. (Gramineae), 5, 204, 255, 256, 257, 275, 295 Heterococcus pulverarius, 191 Misericoccus arenarius, 204 Phenacoccus graminosus, 255 Phenacoccus solani, 273 Pseudococcus fragilis, 295 Rhizoecus cacticans, 379 Lolium perenne (Gramineae), 191, 193, 381 Lophophora williamsi (Cactaceae), 418 Spilococcus cactearum, 416 loquat. See Eriobotrya japonica Lotus sp. (Leguminosae), 5, 219, 221, 225, 227, 253, 262, 263, 264, 318, 389, 391 Phenacoccus artemisiae, 219 Phenacoccus celtisifoliae, 225 Phenacoccus eriogoni, 242 Phenacoccus gossypii, 248 Phenacoccus lotearum, 262 Phenacoccus solani, 273 Pseudococcus fragilis, 295 Pseudococcus obscurus, 313 Puto californicus, 339 Rhizoecus falcifer, 389 lupine. See Lupinus Lupinus sp. (Leguminosae), 231, 234, 238, 277, 279 Chnaurococcus trifolii, 83 Phenacoccus defectus, 231 Phenacoccus echeveriae, 238 Phenacoccus solenopsis, 277 Lupinus albifrons (Leguminosae), 84, 85

HOST Lycium sp. (Solanaceae), 264, 265 Phenacoccus lycii, 264 Lycopersicon sp. (Solanaceae), 252, 254, 275, 276 Ferrisia virgata, 179 Phenacoccus gossypii, 248 Phenacoccus solani, 273 Pseudococcus obscurus, 313 Lycopersicon exculentum (Solanaceae), 181 Lycopersicon esculentum var. commune (Solanaceae), 253, 315 Madura sp. (Moraceae), 181 Ferrisia virgata, 179 Madia gracilis (Compositae), 322, 324 Pseudococcus sorghiellus, 322 Magnolia sp. (Magnoliaceae), 181, 282, 298, 315, 318 Ferrisia virgata, 179 Planococcus citri, 280 Pseudococcus fragilis, 295 Pseudococcus obscurus, 313 Mahonia sp. (Berberidaceae), 227 Phenacoccus colemani, 227 Pseudococcus fragilis, 295 Mahonia nervosa (Berberidaceae), 297, 298 maidenhair fern. See Adiantum Malacothrix sp. (Compositae), 253 Phenacoccus eschscholtziae, 244 Phenacoccus gossypii, 248 Spilococcus eriogoni, 423 Malacothrix californica var. glabrata (Compositae), 5 Malacothrix glabrata (Compositae), 244, 247 Malacothrix saxatilis var. tenuifolia (Compositae), 247 Malacothrix sonchoides (Compositae), 425 mallow. See Malva mallow family. See Malvaceae Malus sp. (Rosaceae), 303 Planococcus citri, 280 Pseudococcus fragilis, 295 Pseudococcus longispinus, 303 Pseudococcus maritimus, 306 Pseudococcus obscurus, 313 Malus pumila (Rosaceae), 315

Malus sylvestris (Rosaceae), 8, 282, 295, 306, 308, 315, 317, 318 Malva sp. (Malvaceae), 254, 275, 276, 295 Phenacoccus gossypii, 248 Phenacoccus solani, 273 Pseudococcus fragilis, 295 Malva rotundifolia (Malvaceae), 277 Malvaceae, 276 Mammillaria sp. (Cactaceae), 316, 317, 381, 418, 419 Ferrisia virgata, 179 Pseudococcus obscurus, 313 Rhizoecus cacticans, 379 Spilococcus cactearum, 416 Mammillaria dolichocentron (Cactaceae), 418 Mammillaria leydeii (Cactaceae), 182 Mammillaria macdougalii (Cactaceae), 418 Mangifera sp. (Anacardiaceae), 303 Planococcus citri, 280 Pseudococcus longispinus, 303 Mangifera indica (Anacardiaceae), 282 mango. See Mangifera mangrove. See Rhizophora manzanita. See Arctostaphylos maple. See Acer Maranta conspicua (Marantaceae), 317 Pseudococcus obscurus, 313 marigold. See Tagetes Marrubium sp. (Labiatae), 254 Phenacoccus eremicus, 240 Phenacoccus gossypii, 248 Marrubium vulgare (Labiatae), 240, 242, 254 matchweed. See Gutierrezia sarothrae Matthiola sp. (Cruciferae), 391, 431 Rhizoecus falcifer, 389 Spilococcus implicatus, 431 maul oak. See Quercus chrysolepsis Medicago sativa (Leguminosae), 5, 9, 12, 254, 255, 283, 315, 317, 396, 397, 398 Phenacoccus gossypii, 248 Phenacoccus graminosus, 255 Planococcus citri, 280 Pseudococcus obscurus, 313 Rhizoecus kondonis, 384 Melilotus sp. (Leguminosae), 295 Pseudococcus fragilis, 295 Mentzelia sp. (Loasaceae), 188

Heliococcus stachyos, 186 Merismostigma astrophytum (Rubiaceae), 105 Chorizococcus lounsburyi, 103 Mesembryanthemum sp. (Aiozoaceae), 306, 308, 315, 317, 347, 350, 363, 365, 366 Pseudococcus maritimus, 306 Pseudococcus obscurus, 313 Puto echinatus, 347 Puto yuccae, 363 mesquite. See Prosopis Mexican orange. See Choisya Mexican tea, joint pine. See Ephedra milk vetch. See Astragalus milkweed. See Asclepias milkweed family. See Asclepiadaceae milkwort. See Polygalä mistletoe. See Phoradendron mock orange. See Philadelphus Monanthocloe litteralis (Gramineae), 154 Distichlicoccus salinus, 154 Monardella sp. (Labiatae), 186, 231,234 Chorizococcus fistulosus, 99 Heliococcus stachyos, 186 Phenacoccus defectus, 231 Monardella odoratissima (Labiatae), 99, 100, 101 monkey flower. See Diplacus ( = Mimulus) Monstera sp. (Araceae), 210 Nipaecoccus nipae, 208 Monterey cypress. See Cupressus macrocarpa Monterey pine. See Pinus radiata Montia perfoliata (Portulacaceae), 113, 115 Chorizococcus rostellum, 113 Morus sp. (Moraceae), 181, 210, 306 Ferrisia virgata, 179 Nipaecoccus nipae, 208 Pseudococcus maritimus, 306 mountain misery. See Chamaebatia foliolosa mulberry. See Morus Musa sp. (Musaceae), 160, 162, 282, 317 Dysmicoccus brevipes, 160 Planococcus citri, 280 Pseudococcus obscurus, 313 muskmelon. See Cucumis Myoporum sp. (Myoporaceae), 303 Pseudococcus longispinus, 303

INDEX

511

Myrsine africana (Myrsinaceae), 318 Pseudococcus obscurus, 313 Myrtillocactus sp. (Cactaceae), 418 Chorizococcus brevicruris, 93 Spilococcus cactearum, 416 Myrtillocactus geometrizans (Cactaceae), 7, 93, 95, 418 myrtle. See Myrtus Myrtus compacta (Myrtaceae), 317 Pseudococcus obscurus, 313 Nama demissum (Hydrophyllaceae), 244, 247 Phenacoccus eschscholtziae, 244 Nandina domestica (Berberidaceae), 317 Pseudococcus obscurus, 313 Narcissus sp. (Amaryllidaceae), 103, 275, 306, 308, 317 Chorizococcus lounsburyi, 103 Phenacoccus solani, 273 Pseudococcus maritimus, 306 Pseudococcus obscurus, 313 nasturtium. See Tropaeolum Nasturtium sp. ( = Tropaeolum, of gardeners) (Tropaeolaceae), 389, 391 Rhizoecus falcifer, 389 Natal grass. See Tricholaena repens needle grass. See Aristida Neocardenasia arquipensis? (Cactaceae), 182 Ferrisia virgata, 179 Nephelium litchi ( = Litchi chinensis) (Sapindaceae), 181 Ferrisia virgata, 179 Nephrolepis sp. (Polypodiaceae, 303 Pseudococcus longispinus, 303 Nephrolepis bostoniensis (Polypodiaceae), 305 Nephthytis sp. (Araceae), 181, 182

Ferrisia virgata, 179 Nerine sp. (Amaryllidaceae), 105 Chorizococcus lounsburyi, 103 Nerine flexuosa (Amaryllidaceae), 103 Nerium sp. (Apocynaceae), 181, 303, 397, 398 Ferrisia virgata, 179 Planococcus citri, 280 Pseudococcus longispinus, 303

512

HOST

INDEX

Nerium sp. (Cont.) Pseudococcus obscurus, 313 Rhizoecus kondonis, 394 Rhizoecus leucosomus, 398 Spilococcus pressus, 442 Nerium oleander (Apocynaceae),283, 305, 315, 318, 398, 442, 444 nettle. See Urtica New Zealand flax. See Phormium tenax Nicotiana sp. (Solanaceae), 282 Planococcus citri, 280 nievitas. See Cryptantha nightshade. See Solanum Nipa fruticans (Palmaceae), 210 Nipaecoccus nipae, 208 Norfolk Island pine. See Araucaria excelsa Norway maple. See Acer platanoides Notholcus lanatus (Gramineae), 172 Dysmicoccus timberlakei, 172 nursery, 6, 7, 9, 10, 12, 16 nut grass, 113, 276. See Cyperus rotundus oak. See Quercus oat grass. See Danthonia californica oats. See Avena Ocotea catesbyana (Lauraceae), 81 Cataenococcus olivaceus, 81 ocotillo. See Fouquieria splendens Odontoglossum sp. (Orchidaceae), 311, 313 Pseudococcus importatus, 298 Pseudococcus microcirculus, 311 Pseudococcus obscurus, 313 Odontoglossum grande (Orchidaceae), 298, 300, 316 Oenothera sp. (Onagraceae), 363, 365, 442, 444 Phenacoccus gossypii, 248 Puto echinatus, 347 Puto yuccae, 363 Spilococcus pressus, 442 Oenothera cheiranthifolia (Onagraceae), 253, 347 Olea sp. (Oleaceae), 181 Ferrisia virgata, 179 Planococcus kraunhiae, 283 Olea chrysophylla (Oleaceae), 283 Olea europaea (Oleaceae), 283 oleander. See Nerium olive. See Olea Onagraceae, 227, 230

Oncidium sp. (Orchidaceae), 298, 300, 313 Pseudococcus importatus, 298 Pseudococcus microcirculus, 311 one-leaved pinyon pine. See Pinus monophylla Ophiopogon sp. (Liliaceae), 389, 391 Rhizoecus falcifer, 389 Opuntia sp. (Cactaceae), 314, 315, 316, 317 Pseudococcus obscurus, 313 orange, 253, 282, 283, 295, 296, 297, 305, 316, 365, 394. See Citrus sinensis orchid (Orchidaceae), 7, 160, 276, 298, 299, 311, 312, 313. See several genera Orchidaceae, 298, 303 orchid family. See Orchidaceae Oriental arbor-vitae. See Thuja orientalis ornamental, 6, 7, 8 Ornithogalum sp. (Liliaceae), 105 Chorizococcus lounsburyi, 103 Orobanche sp. (Orobanchaceae), 275, 277 Phenacoccus solani, 273 Phenacoccus solenopsis, 277 Pseudococcus longisetosus, 300 Orobanche tuberosa (Orobanchaceae), 302 orpine family. See Crassulaceae osage orange. See Maclura Oryza latifolia (Gramineae), 282 Planococcus citri, 280 Oryzopsis hymenoides (Gramineae), 150, 152 Distichlicoccus californicus, 150 Our Lord's Candle. See Yucca whipplei painted cup. See Castilleia palm, 317, 318, 389, 391. See several genera palmetto palm. See Sabal Pandanus sp. (Pandanaceae), 305 Pseudococcus longispinus, 303 Pandanus utilis (Pandanaceae), 305 panic grass. See Panicum Panicum sp. (Gramineae), 160 Antonina graminis, 76 Dysmicoccus brevipes, 160 Panicum purpurascens (Gramineae), 76 paragrass. See Panicum purpurascens

Parasela sp. ( = Dalea) emoryi (Leguminosae), 426 Phenacoccus helianthi, 257 Spilococcus eriogoni, 423 Parosela spinosa (Leguminosae), 259 Parthenocissus inserta (Vitaceae), 297 Pseudococcus fragilis, 295 Parthenocissus quinquefolia (Vitaceae), 255 Phenacoccus gossypii, 248 partridge pea. See Chamaecrista Pasania sp. ( = Lithocarpus) (Fagaceae), 317 Anisococcus quercus, 67 Dysmicoccus quercicolus, 166 Pseudococcus obscurus, 313 Pasania densiflora (Fagaceae), 70, 166 Passiflora sp. (Passifloraceae), 315, 317, 318 Pseudococcus longispinus, 303 Pseudococcus obscurus, 313 Passiflora edulis (Passifloraceae), 305 passion flower. See Passiflora pasture, 8 pea or pulse family. See Leguminosae peach. See Prunus persica pearly everlasting. See Anaphalis margaritacea pear, 6, 12. See also Pyrus Pedicularis densiflora ( Scrophulariaceae ), 227, 230 Phenacoccus colemani, 227 Pelargonium sp. (Geraniaceae), 254, 255, 389, 391 Phenacoccus gossypii, 248 Rhizoecus falcifer, 389 Rhizoecus kondonis, 394 Pelargonium inquinans (Geraniaceae), 397 Peltiphyllum peltatum (Saxifragaceae), 305 Pseudococcus longispinus, 303 Pennisetum purpureum (Gramineae), 160 Dysmicoccus brevipes, 160 Penstemon sp. (Scrophulariaceae), 276 Phenacoccus solani, 273 pepper. See Piper perennial eriastrum, see Eriastrum densifolium Pereskiopsis diqueti (Cactaceae), 316 Pseudococcus obscurus, 313 periwinkle. See Vinca

Persea sp. (Lauraceae), 210, 297, 298, 303, 305 Nipaecoccus nipae, 208 Planococcus citri, 280 Pseudococcus fragilis, 295 Pseudococcus longispinus, 303 Pseudococcus obscurus, 313 Persea americana (Lauraceae), 282, 315 Persea gratissima (Lauraceae), 316 persimmon. See Diospyros Petalonyx sp. (Loasaceae), 219, 240 Phenacoccus alleni, 216 Phenacoccus eremicus, 240 Phenacoccus gossypii, 248 Phenacoccus solenopsis, 277 Puto mimicus, 352 Petalonyx thurberi (Loasaceae), 216, 219, 253, 279, 352, 354 Peteria sp. (Leguminosae), 254 Phenacoccus gossypii, 248 Petunia sp. (Solanaceae), 389 Rhizoecus falcifer, 389 Peucephyllum sp. (Compositae), 242, 266 Phenacoccus eriogoni, 242 Phenacoccus megaulus, 266 Phacelia sp. (Hydrophyllaceae), 227, 230, 231, 234, 276, 279, 442, 444 Phenacoccus colemani, 227 Phenacoccus defectus, 231 Phenacoccus solani, 273 Phenacoccus solenopsis, 277 Spilococcus pressus, 442 Phacelia ramosissima ( Hydrophy llaceae ), 242, 244 Phalaris tuberosa (Gramineae), 8, 255 Phenacoccus graminosus, 255 Phaseolus sp. (Leguminosae), 181, 254 Ferrisia virgata, 179 Phenacoccus gossypii, 248 Phaseolus limensis (Leguminosae), 252, 282 Philadelphus sp. (Saxifragaceae), 297 Pseudococcus fragilis, 295 Philodendron sp. (Araceae), 182, 211, 254, 283 Ferrisia virgata, 179 Nipaecoccus nipae, 208 Phenacoccus gossypii, 248 Planococcus citri, 280 Phleum pratense (Gramineae), 191, 193, 398 Heterococcus pulverarius, 191 Rhizoecus leucosomus, 398

HOST Phoenix sp. (Palmaceae), 211 Nipaecoccus nipae, 208 Planococcus citri, 280 Pseudococcus fragilis, 295 Rhizoecus falcifer, 389 Phoenix canariensis (Palmaceae), 211, 391 Phoenix dactylifera (Palmaceae), 282, 297 Phoenix robelenii (Palmaceae), 389, 391 Phoradendron sp. (Loranthaceae), 315, 318, 446 Pseudococcus obscurus, 313 Spilococcus prosopidis, 444 Phormium sp. (Agavaceae), 389, 465, 467 Pseudococcus longispinus, 303 Rhizoecus falcifer, 389 Trionymus diminutus, 465 Phormium tenax (Agavaceae), 305, 465, 466, 467 Photinia arbutifolia (Rosaceae), 5, 225, 226, 227 Phenacoccus celtisifoliae, 225 Phyllocactus sp. (Cactaceae), 381 Rhizoecus cacticans, 379 Phyllodoce ( = Bryanthus) breweri (Ericaceae), 339, 340 Puto californicus, 339 Phyllostachys bambusoides ( = P. quilioi) (Gramineae), 74 Antonina crawii, 74 Phyllostachys nigra (Gramineae), 74 Picea sp. (Pinaceae), 389, 391 Phenacoccus minimus, 268 Pseudococcus obscurus, 313 Pseudococcus sparsus, 324 Puto sandini, 342 Puto cupressi, 343 Rhizoecus falcifer, 389 Picea breweriana (Pinaceae), 268, 269 Picea engelmanni (Pinaceae), 18, 328, 342, 343 Picea pungens (Pinaceae), 268 Picea pungens glauca (Pinaceae), 316 pickleweed. See Salicornia ambigua pineapple, 10. See also Ananas comosus pineapple family. See Bromeliaceae pink. See Dianthus pine. See Pinus Pinus sp. (Pinaceae), 127, 128, 129, 282, 318 Crisicoccus pini, 127

Dysmicoccus aciculus, 158 Dysmicoccus pinicolus, 164 Phenacoccus helianthi, 257 Planococcus citri, 280 Pseudococcus longispinus, 303 Pseudococcus obscurus, 313 Pseudococcus sparsus, 324 Puto cupressi, 343 Puto laticribellum, 350 Puto pricei, 357 Pinus albicaulis (Pinaceae), 343, 357, 359 Pinus contorta var. latifolia (Pinaceae). 357. 358. 359 Pinus halepensis? (Pinaceae), 129 Pinus jeffreyi (Pinaceae), 329, 350, 352 Pinus macrocarpa (Pinaceae), 129 Pinus monophylla (Pinaceae), 164 Pinus palustris (Pinaceae), 282 Pinus pentaphylla (Pinaceae), 127 Pinus pinea (Pinaceae), 305 Pinus ponderosa (Pinaceae), 18, 259, 328, 343, 350, 351, 352 Pinus radiata (Pinaceae), 127, 129, 158, 159, 315, 317, 343 Pinus thunbergii (Pinaceae), 127 Pinus torreyana (Pinaceae), 343 Piper sp. (Piperaceae), 389 Rhizoecus falcifer, 389 Piptanthocereus sp. (Cactaceae), 418 Spilococcus cactearum, 416 Pittosporum sp. (Pittosporaceae), 297, 305, 315, 316, 317 Pseudococcus fragilis, 295 Pseudococcus longispinus, 303 Pseudococcus obscurus, 313 Pittosporum tobira (Pittosporaceae), 5, 297 Pittosporum undulatum (Pittosporaceae), 297, 316 plane tree. See Platanus Plantago sp. (Plantaginaceae), 244, 247, 276 Phenacoccus eschscholtziae, 244 Phenacoccus solani, 273 Puto pacificus, 354 Plantago juncoides (Plantaginaceae), 354, 356, 357

Plantago lanceolata (Plantaginaceae), 276 plantain. See Plantago Plantanus acerifolia (Platanaceae), 317 Plantanus orientalis (Platanaceae), 283 Platanus sp. (Platanaceae), 113, 115, 291, 293, 298, 317 Chorizococcus rostellum, 113 Planococcus kraunhiae, 283 Pseudococcus dispar, 291 Pseudococcus fragilis, 295 Pseudococcus obscurus, 313 Platanus racemosa (Platanaceae), 315, 318 Pleioblastus ( = A r u n d i n a ria) simonii (Gramineae), 74 Antonina crawii, 74 Pluchea sp. (Compositae), 259 Phenacoccus eremicus, 240 Phenacoccus eriogoni, 242 Phenacoccus helianthi, 257 Spilococcus atriplicis, 414 Pluchea sericea (Compositae), 240, 242, 244, 259 416 Poa sp. (Gramineae), 5, 140, 191, 192, 193 Discococcus graminis, 140 Heterococcus arenae, 189 Heterococcus pulverarius, 191 Tridiscus multiorbis, 457 Poa douglasii (Gramineae), 189, 190, 191 Poa sandbergii (Gramineae), 457, 458, 459 Podocarpus sp. (Podocarpaceae), 317 Pseudococcus obscurus, 313 poinsettia. See Euphorbia pulcherrima poison hemlock. See Conium maculatum poison ivy. See Toxicodendron ( = R h u s ) poison oak. See Rhus diversiloba Poly gala capitata (Polygalaceae), 400 Rhizoecus pritchardi, 400 Polygala cornuta (Polygalaceae), 324 Pseudococcus sorghiellus?, 322 Polygonum sp. (Polygonaceae), 275, 277, 381, 383 Phenacoccus solani, 273 Rhizoecus californicus, 381 pomegranate. See Punica

INDEX

513

poplar, aspen. See Populus popper plant. See Ruellia malacosperma Populus sp. (Salicaceae), 186, 291, 292, 293 Heliococcus stachyos, 186 Phenacoccus minimus, 268 Planococcus citri, 280 Pseudococcus dispar, 291 Populus tremuloides (Salicaceae), 268 Populus trichocarpa (Salicaceae), 283 Porlieria angustifolia (Zygophyllaceae), 81 Cataenococcus olivaceus, 81 Portulaca sp. (Portulacaceae), 160. 275, 277 Dysmicoccus brevipes, 160 Phenacoccus solani, 273 Rhizoecus kondonis, 394 Portulaca grandiflora (Portulacaceae), 397 Portulaca oleracea (Portulacaceae), 276, 277 potato, 13. See also Solanum tuberosum prickly pear. See Opuntia primrose. See Primula Primula sp. (Primulaceae), 305, 316 Phenacoccus gossypii, 248 Pseudococcus longispinus, 303 Pseudococcus obscurus, 313 Primula obconica (Primulaceae), 253 privet. See Ligustrum Prosopis sp. (Leguminosae), 259, 436?, 444, 445, 446 Ferrisia virgata, 179 Phenacoccus helianthi, 257 Spilococcus larreae, 436 Spilococcus prosopidis, 444 Prosopis juliflora (Leguminosae), 181 prune. See Prunus domestica Prunus sp. (Rosaceae), 186, 289, 295, 306, 318, 389, 397, 398 Heliococcus stachyos, 186 Phenacoccus dearnessi, 231 Phenacoccus gossypii, 248 Pseudococcus fragilis, 295 Pseudococcus maritimus, 306 Pseudococcus obscurus, 313 Rhizoecus falcifer, 389 Rhizoecus kondonis, 394 Prunus amygdalus (Rosaceae), 308

514

HOST

INDEX

Prunus armeniaca (Rosaceae), 317 Prunus domestica (Rosaceae), 5 Prunus havardii (Rosaceae), 231 Prunus persica (Rosaceae), 254, 297, 316,389 Prunus persica nectarina (Rosaceae), 317 Prunus serotina (Rosaceae), 298 Pseudotsuga taxifolia (Pinaceae), 343, 360, 361 Puto cupressi, 343 Puto profusus, 360 Psidium sp. (Myrtaceae), 181, 210 Ferrisia virgata, 179 Nipaecoccus nipae, 208 Planococcus citri, 280 Psidium guava (Myrtaceae), 282 Psoralea macrostachya (Leguminosae), 306, 308 Chorizococcus psoraleae, 109 Pseudococcus maritimus, 306 Rhizoecus leucosomus, 398 Psoralea physodes (Leguminosae), 109, 110, 111 Psoralea tenuiflora (Leguminosae), 398 Pteryxia petraea (Umbelliferae), 270, 271 Phenacoccus multisetosus, 268 Ptiloria sp. (=Stephanomeria (Compositae), 259 Phenacoccus helianthi, 257 Ptychosperma elegans (Palmaceae), 389 Rhizoecus falcifer, 389 Pueraria sp. (Leguminosae), 254 Phenacoccus gossypii, 248 pumpkins, squashes. See Cucurbita puncture weed. See Tribulus terrestris Punica sp. (Punicaceae), 298 Planococcus citri, 280 Pseudococcus fragilis, 295 Punica granatum (Punicaceae), 282, 297, 298 Purshia glandulosa (Rosaceae), 242 Phenacoccus eriogoni, 242 Purshia tridentata (Rosaceae), 5, 70 Anisococcus quercus, 67 purslane. See Portulaca Pyracantha sp. (Rosaceae), 254, 297 Phenacoccus gossypii, 248 Pseudococcus fragilis, 295

Pyrus communis (Rosaceae), 282, 295, 306, 308, 309, 315, 316, 317, 318 Planococcus citri, 280 Pseudococcus fragilis, 295 Pseudococcus maritimus, 306 Pseudococcus obscurus, 313 Pyrus malus (=Malus sylvestris) (Rosaceae), 282, 317 Planococcus citri, 280 Pseudococcus obscurus, 313 quackgrass. See Agropyron repens Quercus sp. (Fagaceae), 166, 446, 447, 448 Anisococcus quercus, 67 Chnaurococcus trifolii, 83 Chnaurococcus villosa, 86 Dysmicoccus quercicolus, 166 Pseudococcus obscurus, 313 Puto yuccae, 363 Rhizoecus bituberculatus, 376 Rhizoecus browni, 376 Rhizoecus spinosus, 404 Spilococcus quercinus, 446 Quercus agrifolia (Fagaceae), 85, 86, 87, 166, 168, 315, 318, 404, 405 Quercus chrysolepsis (Fagaceae) 69, 70, 166, 167, 168 Quercus dumosa (Fagaceae), 365 Quercus kelloggi (Fagaceae), 404, 405 Quercus wislizenii (Fagaceae), 376, 377, 379 quince. See Cydonia rabbitbrush. See Chrysothamnus radish. See Raphanus ragweed. See Ambrosia Ramona (=Salvia) polystachya (Labiatae), 61, 363 Anisococcus crawii, 59 Puto yuccae, 363 Ramona (=Salvia) stachyoides (Labiatae), 306, 308, 366 Pseudococcus maritimus, 306 Puto yuccae, 363 Raphanus sativus (Cruciferae), 275, 276 Phenacoccus solani, 273 raspberry. See Rubus rattle box. See Crotalaria red bilberry. See Vaccinium parvifolium

redbud, Judas tree. See Cercis red heather. See Phyllodoce (=Bryanthus) breweri red pepper. See Capsicum redroot. See Lachnanthes redwood. See Sequoia sempervirens redwood trees, 18 Rheum rhaponticum (Polygonaceae), 295, 298 Pseudococcus fragilis, 295 Rhizophora sp. (Rhizophoraceae), 81 Cataenococcus olivaceus, 81 Rhodesgrass, 7, 8. See also Chloris gayana Rhododendron sp. (Ericaceae), 295, 305 Pseudococcus fragilis, 295 Pseudococcus longispinus, 303 Rhoeidium mirophyllum (Anacardiaceae), 181 Ferrisia virgata, 179 rhubarb. See Rheum Rhus diversiloba (Anacardiaceae), 186, 188, 301, 302, 306, 308 Heliococcus stachyos, 186 Pseudococcus longisetosus, 300 Pseudococcus maritimus, 306 Ribes sp. (Saxifragaceae), 389 Pseudococcus fragilis, 295 Rhizoecus falcifer, 389 Ribes sanguineum (Saxifragaceae), 295 rice. See Oryza rice grass. See Oryzopsis rock, 5, 18, 19, 23 rock rose. See Cistus Roebelia sp. (Palmaceae), 161, 162

Dysmicoccus brevipes, 160 roots, 5, 9, 16, 18, 19 Rosa sp. (Rosaceae), 282, 295, 297, 397 Planococcus citri, 280 Pseudococcus fragilis, 295 Rhizoecus kondonis, 394 Rosaceae, 67 rose. See Rosa rose bay. See Rhododendron rose family. See Rosaceae rose mallow. See Hibiscus rose of Sharon, 254, 255. See also Hibiscus syriacus Rubus sp. (Rosaceae), 227, 228, 230, 295, 306, 308, 322, 323 Phenacoccus colemani, 227 Pseudococcus fragilis, 295 Pseudococcus maritimus, 306

Pseudococcus sorghiellus, 322 Rubus vitifolius (Rosaceae), 227, 230, 306, 308 Rudbeckia sp. (Compositae), 254 Phenacoccus gossypii, 248 Ruellia malacosperma (Acanthaceae), 7, 181 Ferrisia virgata, 179 Rumex sp. (Polygonaceae), 275, 276, 397 Phenacoccus eschscholtziae, 244 Phenacoccus solani, 273 Rhizoecus kondonis, 394 Rumex hymenosepalus (Polygonaceae), 244, 247 Russian thistle. See Salsola rye. See Secale cereale ryegrass, 8. See also Lolium Sabal umbraculifera (Palmaceae), 389 Rhizoecus falcifer, 389 sage, 365. See also Salvia sagebrush. See Artemisia sago palm. See Cycas revoluta salal. See Gaultheria shallon salt grass, 454, 455, 456. See also Distichlis St. Augustine grass, 7. See also Stenotaphrum secundatum Saintpaulia sp. (Gesneriaceae), 9, 10, 14, 250, 282, 316, 400, 402 Phenacoccus gossypii, 248 Planococcus citri, 280 Pseudococcus obscurus, 313 Rhizoecus cacticans, 379 Rhizoecus pritchardi, 400 Saintpaulia ionantha (Gesne riaceae), 381 401, 402 Salicornia ambigua (Chenopodiaceae), 315, 317, 333, 334, 335 Pseudoccus obscurus, 313 Puto ambiguus, 333 Salix sp. (Salicaceae), 315 Pseudococcus obscurus, 313 Salsola kali (Chenopodiaceae), 276, 347 Phenacoccus solani, 273 Puto echinatus, 347 Salsola kali var. tenuifolia (Chenopodiaceae), 347, 350 salt grass. See Distichlis saltbush. See Atriplex saltbush family. See Chenopodiaceae saltbrush. See Atriplex lentiformis salt marshes, 16 saltwort. See Batis Salvia sp. (Labiatae), 253, 397

HOST Salvia sp. (Cont.) Amonostherium lichtensioides, 52 Anisococcus crawii, 59 Anisococcus didymus, 61 Phenacoccus eschscholtziae, 244 Phenacoccus gossypii, 248 Phenacoccus helianthi, 257 Puto yuccae, 363 Salvia apiana (Labiatae), 60, 61 Salvia carnosa (Labiatae), 61, 259, 260 Salvia columbariae (Labiatae), 246, 247, 366 Salvia polystachya (Labiatae), 363 Sambucus caerulea (Caprifoliaceae), 315, 316,318 Pseudococcus obscurus, 313 Sambucus glauca (Caprifoliaceae),315, 316, 318 Pseudococcus obscurus, 313 sandbur. See Cenchrus sand dunes, 19 sandpaper plant. See Petalonyx sand verbena. See Abronia sandwort. See Arenaria verna saw- or scrub palmetto. See Serenoa Scabiosa sp. (Dipsaceae), 397, 404, 405 Rhizoecus kondonis, 394 Rhizoecus spinosus, 404 scabious, mourning bride, pincushion flower. See Scabiosa Scalybroom. See Lepidospartum Schinus molle (Anacardiaceae), 295 Pseudococcus fragilis, 295 Sciaphyllum sp. (Araliaceae), 282 Planococcus citri, 280 Scilla amoera (Liliaceae), 105 Chorizococcus lounsburyi, 103 Scotch broom, 317. See also Cytisus scoparius scratch bush. See Urera screw pine. See Pandanus sea blite. See Suaeda Seaforthia elegans (=Ptychosperma elegans) (Palmaceae), 389 Rhizoecus falcifer, 389 seagrape. See Coccoloba sea urchin cacti. See Echinopsis Sebastiania bilocularis (Euphorbiaceae), 363 Puto yuccae, 363 Secale cereale (Gramineae), 485, 487

Trionymus smithii, 485 sedge. See Carex Sedum sp. (Crassulaceae), 276 Phenacoccus solani, 273 Selaginella sp. (Selaginellaceae), 276 Phenacoccus solani, 273 Sempervivum sp. (Crassulaceae), 381 Phenacoccus defectus, 231 Rhizoecus cacticans, 379 Rhizoecus leucosomus, 398 Sempervivum tectorum (Crassulaceae), 231, 234, 398 Senecio mikanioides (Compositae), 254 Phenacoccus gossypii, 248 Senecio péndula (Compositae), 389 Rhizoecus falcifer, 389 senna. See Cassia Sequoia sempervirens (Taxodiaceae), 5, 343, 412, 414, 431, 432, 434, 449, 450 Puto cupressi, 343 Spilococcus andersoni, 412 Spilococcus implicatus, 431 Spilococcus sequoiae, 448 Serenoa sp. (Palmaceae), 303 Pseudococcus longispinus, 303 service berry. See Amelanchier sp. Setaria viridis (Gramineae), 113, 115, 191 Chorizococcus rostellum, 113 Heterococcus pulverarius, 191 sheaths, 5 shooting star. See Dodecatheon shore grass. See Monanthocloe Sida sp. (Malvaceae), 160, 442, 444 Dysmicoccus brevipes, 160 Phenacoccus eremicus, 240 Phenacoccus solani, 273 Spilococcus pressus, 442 Sida hederacea (Malvaceae), 240, 242, 275, 277 silktassel. See Garrya silver spruce. See Picea pungens Simmondsia chinensis (Buxaceae), 360, 362, 363 Puto simmondsiae, 360 Sinningia sp. (Gesneriaceae), 387, 388 Rhizoecus eluminatus, 387 Sitanion sp. (Gramineae), 130

Chorizococcus polyporous, 107 Cryptoripersia salina, 130 Discococcus spectabilis, 144 Rhizoecus browni, 376 Trionymus quadricirculus, 483 Sitanion grass, 376. See also Sitanion Sitanion hystrix (Gramineae), 107, 130, 144, 147, 376, 379, 483, 484, 485 small sedge. See Cyperus tenerrimus snakeweed. See Gutierrezia sneezeweed. See Helenium snowberry. See Symphoricarpos albus soft acanthus. See Acanthus mollis soil, 14, 17, 19, 21 Solanum sp. (Solanaceae) 252, 254, 259, 260, 275, 277, 283, 295, 315, 316, 317, 318 Heliococcus stachyos, 186 Phenacoccus gossypii, 248 Phenacoccus helianthi, 257 Phenacoccus solani, 273 Phenacoccus solenopsis, 277 Planococcus citri, 280 Pseudococcus fragilis, 295 Pseudococcus obscurus, 313 Solanum douglasii (Solanaceae), 253, 259, 260, 283 Solanum melongena (Solanaceae), 253, 282 Solanum pseudocapsicum (Solanaceae), 254 Solanum tuberosum (Solanaceae), 253, 274, 275, 276, 317 Solanum tuberosum sativus (Solanaceae), 276 Solanum umbelliferum (Solanaceae), 186, 188 Solanum xanthi douglasii (Solanaceae), 277 Solidago sp. (Compositae), 5,315,317, 322, 426 Pseudococcus obscurus, 313 Pseudococcus sorghiellus, 322 Spilococcus eriogoni, 423 Sonchus sp. (Compositae), 253 Phenacoccus gossypii, 248 sorghum, 322 Pseudococcus sorghiellus, 322 Sorghum halepense (Gramineae), 7, 76, 113, 115, 487, 489 Antonina graminis 76 Chorizococcus rostellum, 113

INDEX

515

Trionymus utahensis, 487 soursop. See Annona muricata sow thistle. See Sonchus Spanish bayonet, 5. See also Yucca Sparmannia africana (Tiliaceae), 254 Phenacoccus gossypii, 248 spear grass. See Stipa speedwell. See Veronica Sphaeralcea orcutti (Malvaceae), 216, 219 Phenacoccus alleni, 216 spikeweek, 427. See also Centromadia spindle tree. See Euonymus spleenwort. See Asplenium Sprekelia sp. (Amaryllidaceae), 103 Chorizococcus lounsburyi, 103 spring beauty. See Claytonia spruce. See Picea spurge. See Euphorbia Squawcarpet or Mahala Mat. See Ceanothus prostratus squill. See Scilla squirrel tail. See Sitanion Stachys sp. (Labiatae), 188 Heliococcus stachyos, 186 Puto yuccae, 363 Spilococcus pressus, 442 Stachys bullata (Labiatae), 186, 187, 188, 442, 444 Stapelia sp. (Asclepiadaceae) 7, 93, 95 Chorizococcus brevicruris, 93 Stapelia glandiflora (Asclepiadaceae), 93, 95 Stapelia hursuta (Asclepiadaceae), 93 95 Stapelia verrucosa (Asclepiadaceae), 93, 95 star cactus. See Astrophytum star thistle. See Centaurea starwort, Michaelmas daisy. See Aster Stellaria media (Caryophyllaceae), 397 Rhizoecus kondonis, 394 Stenotaphrum secundatum (Gramineae), 76, 77, 78 Stephanomeria sp. (Compositae), 262, 264, 279 Antonina graminis, 76 Phenacoccus helianthi, 257 Phenacoccus lotearum, 262 Phenacoccus megaulus, 266

Phenacoccus solenopsis, 277 Stephanomeria pauciflora (Compositae), 266, 267

516

HOST

INDEX

Stephanomeria virgata (Compositae), 259 Stephanotis sp. (Asclepiadaceae), 282, 389 Planococcus citri, 280 Rhizoecus falcifer, 389 Sterculia urens (Sterculiaceae), 210 Nipaecoccus nipae, 208 Stichwort. See Stellaria Stillingia linearifolia (Euphorbiaceae), 438, 440 Spilococcus parkeri, 438 Stipa sp. (Gramineae), 137, 139, 140, 144, 191, 204, 369, 370 Cryptoripersia salina, 130 Discococcus flabellatus, 137 Discococcus hopi, 142 Distichlicoccus megacirculus, 152 Heterococcus arenae, 189 Misericoccus arenarius, 204 Radicoccus kelloggi, 368 Rhizoecus browni, 376 Stipa lepida (Gramineae), 379 Stipa pulchra (Gramineae), 189, 191 Stipa thurberiana (Gramineae) 130, 142, 143, 152, 153, 154 stock. See Matthiola stonecrop, live-for-ever. See Sedum stone fruits. See Prunus storax. See Styrax storksbill. See Erodium Straussia sp. (Rubiaceae), 160, 210

Dysmicoccus brevipes, 160

Nipaecoccus nipae, 208 strawberry, 9. See also Fragaria Strelitzia sp. (Musaceae), 210, 211, 283, 306, 308 Nipaecoccus nipae, 208 Planococcus citri, 280 Pseudococcus longispinus, 303 Pseudococcus maritimus, 306 Pseudococcus obcurus, 313 Strelitzia reginae (Musaceae), 283, 305, 317 Styrax wilsoni (Styracaceae), 254 Phenacoccus gossypii, 248 Suaeda sp. (Chenopodiaceae), 275, 277, 279, 426, 427 Phenacoccus solani, 273 Phenacoccus solenopsis, 277 Spilococcus eriogoni, 423 Suaeda fruiticosa (Chenopodiaceae), 280 succulent plants, 5, 7

succulents. See several genera sugar apple. See Annona squamosa sulphur-flower. See Eriogonum umbellatum ssp. polyanthum sumac. See Rhus sunflower. See Helianthus sun rose. See Helianthemum sweet clover. See Melilotus sweetfläg. See Acorus sweet orange. See Citrus sinensis sweet potato. See Ipomoea batatas sweet vernal grass. See Anthoxanthum odoratum sword fern. See Nephrolepis Symphoricarpos albus (Caprifoliaceae), 227 Symphoricarpos racemosus (Caprifoliaceae), 227, 230 Phenacoccus colemani, 227 Syntherisma sanguinalis (=Digitaria sanguinalis) (Gramineae), 113, 115 Chorizococcus rostellum, 113 Syringa vulgaris (Oleaceae), 389 Rhizoecus falcifer, 389 Tagetes sp. (Compositae), 253 Phenacoccus gossypii, 248 tall oat grass. See Arrhenatherum elatius tamarisk. See Tamarix Tamarix sp. (Tamaricaceae), 305 Pseudococcus longispinus, 303 tan oak. See Pasania (=Lithocarpus) Tapirira edulis (Anacardiaceae), 306 Pseudococcus maritimus, 306 Taraxacum vulgare (Compositae), 253 Phenacoccus gossypii, 248 tarragon. See Artemisia dracunculus Tarulespinus ferocactus? (Cactaceae), 182 Ferrisia virgata, 179 tarweed. See Hemizonia or Madia Taxaceae, 414, 421, 446 Taxodiaceae, 414, 421, 446 taxodium family. See Taxodiaceae Taxus sp. (Taxaceae), 298, 317, 318 Pseudococcus fragilis, 295 Pseudococcus longispinus, 303 Pseudococcus obscurus, 313 Taxus baccata (Taxaceae), 297

Taxus baccata fastigiata (Taxaceae), 317 Taxus baccata var. washingtonii (Taxaceae), 305 Tecoma jasminoides (Bignoniaceae), 297 Pseudococcus fragilis, 295 teasel. See Dipsacus Tetradymia canescens (Compositae), 271 Phenacoccus multisetosus, 268 Tetradymia spinosa (Compositae), 213, 234 Phenacoccus defectus, 231 Tetraplasandra sp. (Araliaceae), 303 Pseudococcus longispinus, 303 Theobroma cacao (Sterculiaceae), 282 Planococcus citri, 280 Thermopsis sp. (Leguminosae), 186 Heliococcus stachyos, 186 thinleaf huckleberry. See Vaccinium membranaceum thorn apple. See Datura thrift. See Armeria Thuja orientalis (Cupressaceae), 170, 171, 172 Dysmicoccus ryani, 170 thyme. See Thymus Thymus sp. (Labiatae), 389 Rhizoecus falcifer, 389 tibinagua. See Eriogonum nudum tickseed. See Coreopsis tidytips. See Layia Tilia americana (Tiliaceae), 306 Pseudococcus maritimus, 306 Tillandsia punctulata (Bromeliaceae), 162, 163, 164 Dysmicoccus mackenziei, 162 timothy. See Phleum tobacco, 10. See also Nicotiana tomato. See Lycopersicon Torreya californica (Taxaceae), 317, 412, 414 Pseudococcus obscurus, 313 Spilococcus andersoni, 412 touch-me-not, snap weed. See Impatiens torch lily. See Kniphofia Toxicodendron radicans ( = R h u s radicans) (Anacardiaceae), 181 Ferrisia virgata, 179 Tragopogon porrifolius (Compositae), 317 Pseudococcus obscurus, 313 Tribulus cistoides (Zygophyllaceae), 181 Ferrisia virgata, 179

Tribulus terrestris (Zygophyllaceae), 276, 347, 350 Phenacoccus solani, 273 Puto echinatus, 347 Tricholaena repens (Gramineae), 160 Dysmicoccus brevipes, 160 Trifolium sp. (Leguminosae), 322, 324 Phenacoccus graminosus, 255 Pseudococcus sorghiellus, 322 Trifolium fragiferum (Leguminosae), 255 Trifolium repens (Leguminosae), 255 Triticum sp. (Gramineae), 255 Phenacoccus graminosus, 255 Triticum aestivum (Gramineae), 487 Trionymus utahensis, 487 Tritoma sp. (=Kniphofia) (Liliaceae), 105 Chorizococcus lounsburyi, 103 Tropaeolum sp. (Tropaeolaceae), 389, 391 Rhizoecus falcifer, 389 Tropaeolum majus (Tropaeolaceae), 389 tuftroot. See Dieffenbachia turkey mullein. See Eremocarpus setigerus twigs, 16 Ulex europaeus (Leguminosae), 253, 315, 317 Phenacoccus gossypii, 248 Pseudococcus obscurus, 313 Ulmus sp. (Ulmaceae), 186 Heliococcus stachyos, 186 Uniola ornithorhynca (Gramineae), 463 Trionymus caricus, 463 Urera sp. (Urticaceae), 210 Nipaecoccus nipae, 208 Urtica sp. (Urticaceae), 254 Phenacoccus gossypii, 248 Vaccinium sp. (Ericaceae), 134 Cucullococcus vaccinii, 134 Vaccinium membranaceum (Ericaceae), 134, 135 Vaccinium parvifolium (Ericaceae), 134, 135 Vanda sp. (Orchidaceae), 298, 300 Pseudococcus importatus, 298 velvet grass. See Holcus or Notholcus lanatus Verbena sp. (Verbenaceae), 253 Phenacoccus gossypii, 248 Verbesina encelioides var. exauriculata (Compositae), 275, 276

HOST Verbesina encelioides (Com.) Phenacoccus solani, 273 Veronica sp. (Scrophulariaceae), 253, 254, 298, 392 Phenacoccus gossypii, 248 Pseudococcus fragilis, 295 Rhizoecus falcifer, 389 vetchling. See Lathyrus Viburnum sp. (Caprifoliaceae), 254, 318 Phenacoccus gossypii, 248 Pseudococcus obscurus, 313 Viburnum macrocephalum (Caprifoliaceae), 254 Vigna sinensis (Leguminosae), 275 Phenacoccus solani, 273 Vinca sp. (Apocynaceae), 253 Phenacoccus gossypii, 248 Viola sp. (Violaceae), 275 Phenacoccus solani, 273 Spilococcus pressus, 442 Viola sarmentosa (Violaceae), 442, 443, 444 violet. See Viola vipers bugloss. See Echium Virginia creeper. See Ampélopsis quinquefolia (=Parthenocissus quinquefolia) Vitis sp. (Vitaceae), 210, 303, 306, 308, 309, 315, 317, 389

Nipaecoccus nipae, 208 Pseudococcus longispinus, 303 Pseudococcus maritimus, 306 Pseudococcus obscurus, 313 Rhizoecus falcifer, 389 Vitis capensis (Vitaceae), 305 Vriesia sp. (Bromeliaceae), 162, 164 Dysmicoccus mackenziei, 162 Watsonia sp. (Iridaceae), 389, 391 Rhizoecus falcifer, 389 waxberry. See Symphoricarpos wax plant. See Hoya wheat, 8. See also Triticum wheat grass. See Agropyron white fir. See Abies concolor white leaf manzanita. See Arctostaphylos viscida White sage. See Salvia apiana Wilcoxia sp. (Cactaceae), 418 Spilococcus cactearum, 416 wild buckwheat. See Eriogonum wild oats. See Avena fatua wild rye. See Elymus willow. See Salix

wire grass. See Eleusine indica Wisteria sp. (Leguminosae), 253, 282, 285, 297, 316 Phenacoccus gossypii, 248 Planococcus kraunhiae, 283 Pseudococcus fragilis, 295 Pseudococcus obscurus, 313 Wisteria floribunda (Leguminosae), 283 wood betony, lousewort. See Pedicularis wood violet. See Viola sarmentosa Wyethia sp. (Compositae), 253 Phenacoccus gossypii, 248 Xanthium sp. (Compositae), 253 Phenacoccus gossypii, 248 yam. See Dioscorea yarrow. See Achillea yellow star thistle. See Centaurea solstitialis yerba santa. See Eriodictyon yew. See Taxus yew family. See Taxaceae Yucca sp. (Agavaceae), 5, 81, 101, 102, 123, 124, 298, 317, 365, 465, 467

INDEX

517

Cataenococcus olivaceus, 81 Chorizococcus interruptus, 101 Chorizococcus reducta, 111 Chorizococcus rostellum, 113 Chorizococcus snellingi, 117 Chorizococcus yuccae, 123 Pseudococcus fragilis, 295 Pseudococcus obscurus, 313 Puto yuccae, 363 Trionymus diminutus, 465 Yucca brevifolia (Agavaceae), 117, 118, 119 Yucca gloriosa (Agavaceae), 81, 82 Yucca shidigera (Agavaceae), 113, 115, 123 Yucca whipplei (Agavaceae), 363, 364, 365 Zantedeschia sp. (Araceae), 254, 306 Phenacoccus gossypii, 248 Pseudococcus maritimus, 306 Zea mays (Gramineae), 322 Pseudococcus sorghiellus, 322 Zygocactus truncatus (Cactaceae), 315 Pseudococcus obscurus, 313

GENERAL INDEX T h i s index does not include the host plants of mealybugs. T h e host index immediately precedes this section. In the following index the n a m e s listed in italics are synonyms, w h e r e a s those in r o m a n letters represent valid genera or species. P a g e n u m b e r s of the principal references are in bold-face type. M a i n h e a d ings are in capital letters. Both the c o m m o n as well as the scientific n a m e of the mealybugs are listed to help find the desired i n f o r m a t i o n .

Abdomen, 36, 39 aberrans, Pseudococcus, 50, 288, 289, 290, 291, 496 aberrans, Trionymus, 291 Aberrant mealybug. See Pseudococcus aberrans Abnormal mealybug. See Anisococcus abnormalis abnormalis, Anisococcus, 45, 48, 55, 56, 57, 494 Abortive mealybug. See Chorizococcus abortivus abortivus, Chorizococcus, 48, 89, 90, 91, 111, 494 Abronia mealybug. See Chorizococcus abroniae abroniae, Chorizococcus, 48, 88, 91, 92, 93, 95, 494 Acanthomyops claviger californicus, 56 acericola, Phenacoccus, 29, 215 aceris, Phenacoccus, 214 aceris, Pseudococcus, 214 aciculus, Dysmicoccus, 4, 49, 157, 158, 159, 495 acid fuchsin, 34, 35 acirculus, Puto, 5, 50, 327, 329, 330, 332, 497 ACKNOWLEDGMENTS, 2

acritocera, Lachnodiella, 81 Acropyga paramaribensis, 22 adenostomae, Anisococcus, 3, 5, 48, 55, 56, 58, 59, 61, 494 adenostomae, Heliococcus, 49, 183, 185, 495 Pseudococcus, adonidum, 303

Adult female, 24 advena, Phenacoccus, 214, 244, 247 Adversities caused by chemicals, 14 aedaegus, 26 africanus, Rhizoecus, 389 Agave mealybug. See Chorizococcus reducta agavis, Pseudococcus, 289 agrifoliae, Pseudococcus, 86, 166 (misident.) albicans, Puto, 8, 50, 329, 331, 332, 333, 497 Aldrin 13, 14 alkalinus, Distichlicoccus, 148 Allen mealybug. See Phenacoccus alleni alleni, Phenacoccus, 16, 21, 49, 215, 216, 217, 218, 219, 240, 242, 260, 495 altoarcticus, Chorizococcus, 88 ambigua, Puto, 327 Ambiguous mealybug. See Puto ambiguus ambiguus, Puto, 4, 26, 50, 327, 328, 333, 334, 335, 339, 345, 359, 497 American grass mealybug. See Trionymus americanus americanus, Rhizoecus, 372 americanus, Trionymus, 4, 51, 461, 462, 498 Amonostherium, 41, 42, 43, 46, 51, 52, 201, 206 lichtensioides, 3, 16, 48, 52, 53, 54, 494 518

anal lobes, 39, 40 anal lobe seta, 39 anal opening, 39 anal ring, 25, 39, 40 Anarhopus sydneyensis, 7 Anderson mealybug. See Spilococcus andersoni andersoni, Spilococcus, 4, 51, 170, 4 1 2 , 4 1 3 , 4 9 8 Anisococcus, 45, 55, 56 abnormalis, 45, 48, 55, 56, 57, 494 adenostomae, 3, 5, 48, 55, 56, 58, 59, 61, 494 crawii, 3, 48, 55, 59, 60, 61, 494 didymus, 3, 48, 55, 59, 61, 62, 63, 494 ephedrae, 3, 48, 55, 63, 64, 65, 494 imperialis, 48, 55, 65, 66, 67, 494 oregonensis, 4, 5, 48, 55, 67, 68, 70, 494 quercus, 4, 5, 48, 55, 56, 67, 69, 70, 71, 494 Annual-flowering plants (grasses excluded), 17 ANNUAL SUCCESSIONS, 2 4 , 2 7 ANT ASSOCIATIONS, 1 6 , 2 0

Ant constructions, 20 ant tent, 20, 21 Ant transport, 22 antennae 25, 36, 40 antennata, Puto ( = Putonia), 326 Anthelococcus, 47, 71 simondsi, 48, 71, 72, 73, 494 Antonina, 18, 25, 38, 41, 44, 73, 74, 78

crawii, 4, 48, 74, 75, 78, 80, 494 graminis, 4, 7, 8, 10, 13, 18, 29, 48, 74, 76, 77, 80, 494 pretiosa, 4, 48, 74, 76, 78, 79, 80, 494 Antonina group, 41 Antoninoides, 25, 45 ants, 5, 9, 20, 85, 89, 99, 107, 109, 117, 130, 132, 168, 174, 175, 199, 204, 221, 224, 225, 227, 230, 240, 247, 254, 262, 275, 276, 277, 291, 298, 300, 309, 310, 398 aphyllonis, Chorizococcus, 88, 93, 119, 123 apodus, 25 apterous, 26 arctostaphyli, Puto, 4, 26, 50, 329, 332, 335, 336, 337, 497 arenae, Heterococcus, 4, 18, 49, 188, 189, 190, 191, 495 arenaria, Ripersia, 203, 204 arenarius, Misericoccus, 4, 19, 42, 49, 204, 205, 495 arizonensis, Cryptoripersia (misident.), 130 arizonensis. Ripersia, 129 Artemisia mealybug. See Phenacoccus artemisiae artemisiae, Phenacoccus, 4, 16, 17, 20, 49,214,219, 220, 221, 495 artemisiae, Eriococcus, 52 artemisiae, Pseudococcus, 52

GENERAL Arundinaria mealybug. See Pseudantonina arundinariae arundinariae, Pseudantonina, 4, 50, 286, 287, 496 arundinis, Distichlicoccus, 49, 148, 149, 150, 152, 154, 495 Arundo mealybug. See Distichlicoccus arundinis Associated mealybug. See Rhizoecus associatus associatus, Rhizoecus, 50, 372, 373, 374, 497 Atriplex mealybug. See Puto atriplicis atriplicis, Heliococcus, 49, 182, 183, 185, 186, 188, 495 atriplicis, Humococcus, 4, 7, 19, 49, 194, 195, 197, 199, 201, 495 atriplicis, Puto, 50, 328, 337, 338, 339, 497 atriplicis, Spilococcus, 4, 17, 51, 412, 414, 415, 498 Atrococcus, 88 aurantius, Dysmicoccus, 156n aurilanatus, Nipaecoccus, 4, 5, 49, 206, 207, 208, 210, 495 auxiliary setae, 39 Azalea mealybug. See Crisicoccus azaleae azaleae, Crisicoccus, 4, 49, 125, 126, 127, 129, 494 bahiae, Phenacoccus, 363 bakeri, Pseudococcus, 306, 308 bambusae, Antonina, 74 bambusae, Pseudantonina, 285 barbarae, Hypogeococcus, 201, 203 barberi, Puto, 328, 335, 339 bark, 33, 43 Bark mealybug. See Spilococcus corticosus beak, 38 beating sheet, 32 Benefits to ants (by mealybugs), 22 Benefits to mealybugs (by ants), 20 Bergrothia, 459 (preoccupied) Bergrothiella, 459 (preoccupied) Berlese funnel, 33 Bermuda grass mealybug. See Chorizococcus rostellum betheli, Phenacoccus, 231 Betony mealybug. See Heliococcus stachyos bicaudatus, Pseudococcus, 179 bicirculus, Rhizoecus, 50, 371, 374, 375, 497 bi- or tritubular cerores (ducts), 39 Biological control, 14 BIOLOGY, 2 4

Bilberry mealybug. See Cucullococcus vaccinii bispinosus, Dysmicoccus, 157 Bituberculate soil mealybug. See Rhizoecus bituberculatus bituberculatus, Rhizoecus, 50, 371, 376, 377, 404, 497 bitubular, 39 Blind mealybug. See Rhizoecus eluminatus Bluegrass mealybug. See Heterococcus pulverarius body color, 43 body form, 40 body setae, 40 boharti, Rhizoecus, 372 bohemicus, Heliococcus, 182 boninsis, Dysmicoccus, 13, 157 boutelouae, Antonina, 74 brachypodii, Laboulbenia, 73 brachypterous, 26 branches, 33 brevicruris, Chorizococcus, 7, 48, 88, 93, 94, 95, 494 brevipes, Dactylopius, 156 brevipes, Dysmicoccus, 10, 13,24, 29, 49, 157, 160, 161, 162, 495 brevispinus, Dactylopius, 280 bromeliae, Pseudococcus, 160 (misident.) bromi, Trionymus, 487 browni, Rhizoecus, 50, 371, 376, 378, 379, 497 Brown's soil mealybug. See Rhizoecus browni bryanthi, Puto, 4, 50, 328, 335, 339, 342, 359, 497 Bryanthus mealybug. See Puto bryanthi Burning, 14 Burro-weed mealybug. See Spilococcus ventralis cactearum, Spilococcus, 5, 24, 51, 411, 416, 417, 498 Cacticans mealybug. See Rhizoecus cacticans cacticans, Rhizoecus, 4, 5, 50, 371, 374, 379, 380, 381, 497 Cactus mealybug. See Spilococcus cactearum Cajon mealybug. See Phenacoccus cajonensis cajonensis, Phenacoccus, 49, 214, 221, 222, 223, 225, 259, 495 calceolariae, Trionymus, 465 (misident.) calcitectus, Puto, 328, 329 California ground mealybug. See Rhizoecus californicus California nutmeg mealybug. See Puto cupressi

California poppy mealybug. See Phenacoccus eschscholtziae California puto mealybug. See Puto californicus California tea mealybug. See Chorizococcus psoraleae californicus, Chorizococcus 48, 89, 96, 97, 109, 494 californicus, Distichlicoccus, 4, 49, 148, 150, 151, 152, 154, 495 californicus, Puto, 5, 50, 328, 339, 341, 342, 497 californicus, Rhizoecus, 4, 50, 371, 381, 382, 383, 497 californicus, Scaptococcus, 19, 51, 407, 497 campestris, Rhizoecus, 371 Camponotus, 22, 291 Camponotus yogi, 89, 199 Canada balsam, 34 cannae, Pseudococcus, 160 capensis, Phenacoccus, 216 capensis, Pseudococcus, 313, 316 Carex mealybug. See Trionymus caricis caricis, Discococcus, 4, 49, 136, 137, 138, 140, 144, 495 caricis, Trionymus, 4, 51, 460, 463, 498 caritus, Humococcus, 49, 194, 196, 197, 495 carton, 20 casserole, porcelain, 34 casuarinae, Sphaerococcus, 74 Cataenococcus, 46, 80, 81 olivaceus, 4, 5, 48, 81, 82, 83, 494 catalinae, Eriococcus artemisiae variety, 52 ceanothi, Ceroputo yuccae variety, 363 ceanothi, Spilococcus, 51, 412, 419, 420, 498 Celtis mealybug. See Phenacoccus celtisifoliae celtisifoliae, Phenacoccus, 4, 5, 49, 216, 225, 226, 495 cerarii, 25, 39, 40 Cerarii mealybug. See Phenacoccus eriogoni ceraricus, Humococcus, 193 Ceroputo, 214, 326, 333, 363 cevalliae, Phenacoccus, 277 Chaetococcus, 74, 76 Chamise mealybug. See Anisococcus adenostomae Chaparral mealybug. See Spilococcus quercinus chloral-hydrate medium, 34, 35 Chlorinated hydrocarbons, 13 Chnaurococcus, 42, 47, 83

INDEX

519

trifolii, 5, 17, 19, 42, 48, 83, 84 85, 86, 494 villosa, 4, 42, 48, 83, 85, 86, 87, 494 Choosing a location, 32 Chorizococcus, 38, 41, 42, 86, 148, 214 abortivus, 48, 89, 90, 91, 111, 494 abroniae, 48, 88, 91, 92, 93, 95, 494 brevicruris, 7, 48, 88, 93, 94, 95, 494 californicus, 48, 89, 96, 97, 109, 494 coxindex, 48, 88, 96, 98, 99, 494 fistulosus, 48, 88, 99, 100, 101, 494 interruptus, 49, 89, 101, 102, 103, 105, 117, 494 lounsburyi, 5, 49, 89, 101, 103, 104, 105, 494 penultimatus, 49, 89, 105, 106, 107, 494 polyporus, 17, 19, 49, 89, 96, 107, 108, 109, 494 psoraleae, 49, 89, 91, 101, 109, 110, 111, 494 reducta, 4, 49, 88, 111, 112, 113, 123, 494 rostellum, 4, 18, 19, 49, 88, 113, 114, 115, 494 senarius, 49, 89, 115, 116, 117, 494 snellingi, 49, 88, 117, 118, 119, 494 wilkeyi, 49, 86, 119, 120, 121, 494 wilsoni, 49, 88, 99, 101, 121, 123, 494 yuccae, 49, 88, 123, 124, 125, 494 Chromosome number, 30 chromosomes, 28, 29, 30, 31 Circaputo, 45 circulus, 36, 39 citri, Planococcus, 4, 5, 6, 7, 10, 13, 14, 16, 24, 26, 27, 29, 43, 50, 280, 281, 282, 283, 285, 496 citri, Pseudococcus, 280 Citrophilus mealybug. See Pseudococcus fragilis citrophilus, Pseudoccoccus, 295 Citrus mealybug. See Planococcus citri clandestinus, Trionymus, 460 CLASSIFICATION, 3 6 , 4 0

Clavicoccus tribulus, 29 claviger, Trionymus, 461 claviseta, Ferrisia, 179 claw, 26, 38 Clover root mealybug. See Chnaurococcus trifolii Cluster mealybug. See Dysmicoccus racemus clypeus, 38 Coastal grass mealybug. See Tridiscus distichlii Coccidella, 370, 383, 394 Coccophagus gurneyi, 6 Coccus, 83, 280, 322

520

GENERAL

INDEX

cockerellae, Ripersia, 83, 85 cockerelli, Phenacoccus, 231 Coconut mealybug. See Nipaecoccus nipae coffeae, Geococcus, 44n coffeae, Neorhizoecus, 22 Coleman mealybug. See Phenacoccus colemani colemani, Phenacoccus, 4, 5, 19, 23, 49, 216, 225, 227, 228, 229, 230, 234, 248 (misident.), 496 COLLECTING METHODS,

32

Collecting on branches, 33 Collecting on conifers, 33 Collecting on crown and roots, 33 Collecting on foliage and stems, 32 Collecting on grass, 33 comstocki, Pseudococcus, 13, 158 (misident.), 289, 297 confusella, Ripersia, 83 (misident.) confusum, Erium, 52 conical setae, 39, 40 Conifers, 18, 33 CONTROL, 1 2

ceriferus, Dactylopius, 179 CONTROL MEASURES, 1 2

corticosus, Spilococcus, 16, 51, 411, 421, 422, 498 Cottony bamboo mealybug. See Antonina crawii cottony ovisac, 42 Coxa mealybug. See Chorizococcus coxindex coxindex, Chorizococcus, 48, 88, 96, 98, 99, 494 crawii, Anisococcus, 3, 48, 55, 59, 60, 61, 494 crawii, Antonina, 4, 48, 74, 75, 78, 80, 494 crawii, Dactylopius, 55, 363 (as to the male) crawii, Pseudococcus, 67 (misident.) Crematogaster, 20, 22, 117, 221, 227 Crematogaster coarctata, 99 Crematogaster coarctata coarctata, 85, 221, 276 Crematogaster coarctata vermiculata, 224, 225, 247, 262, 276 Crematogaster lineolata, 227, 230, 300 Crisicoccus, 41, 46, 125 azaleae, 4, 49, 125, 126, 127, 129, 494 pini, 4, 18, 49, 125, 127, 128, 129, 494 crop sanitation, 14 crotonis, Pseudococcus, 160 (misident.) crown, 33 crown-infesting mealybugs, 17 Cryptolaemus montrouzieri, 6 Cryptoripersia, 18, 42, 43, 44, 45, 45n, 47, 83, 129, 130

salina, 4, 5, 22, 42, 49, 130, 131, 132, 494 tubulata, 45n, 49, 130, 132, 133, 494 crystalline-like rods, 42 cualatensis, Cataenococcus, 81 Cucullococcus, 46, 134 vaccinii, 4, 49, 134, 135, 136, 494 Cultural methods, 14 cupressi, Ehrhorni, 4, 18, 49, 176, 177, 178, 495 cupressi, Puto, 4, 18, 50, 328, 343, 344, 345, 350, 357, 360, 497 cupressi, Sphaerococcus, 176 cupressicolus, Pseudococcus, 170 cuspidatae, Dysmicoccus, 13, 157, 164, 166 cyperalis, Rhizoecus, 4, 50, 372, 383, 384, 385, 400, 497 Cypress bark mealybug. See Ehrhorni cupressi Cypress mealybug. See Dysmicoccus ryani CYTOLOGY, 2 8 CYTOLOGY ( T E C H N I Q U E ) ,

30

Dactylopius, 52, 59, 63, 67, 81, 83, 125, 127, 154, 160, 170, 179, 206, 208, 280, 283, 303, 306, 322, 343, 363 Dactylopius brevipes, 156 Dactylopius crawii, 55, 363 Dactylopius lichtensioides, 51 Dactylopius nipae, 206 Dactylopius olivaceus, 80 Dactylopius pini, 125 Dactylopius salinus, 147 Dactylopius sequoiae (as to the male), 343 dasychloae, Distichlicoccus, 148 dasychloae, Humococcus, 194 dasylirii, Dactylopius, 179 DDT, 13 dearnessi, Phenacoccus, 4, 49, 214, 231, 232, 496 decorata, Stemmatomerinx, 4, 51, 453, 454, 498 Decorated mealybug. See Puto decorosus decoratus, Rhizoecus, 389 decorosus, Puto, 17, 26, 50, 328, 345, 346, 497 defectus, Phenacoccus, 4, 16, 49, 215, 231, 233, 234, 275, 279, 496 defensive mechanism, 25 DEFINITION (Family Pseudococcidae), 40 denticle, 26, 38 dermal pores, 25 Desert mealybug. See Spilococcus larreae Desert thorn mealybug. See Phenacoccus lycii desertorum, Dysmicoccus, 157

Destitute mealybug. See Phenacoccus destitutus destitutus, Phenacoccus, 49, 214, 235, 236, 264, 496 destructor, Dactylopius, 280 Diazinon, 13 Dichloroethyl ether, 14 Dichlorovos (DDVP), 13 Dicoria mealybug. See Phenacoccus dicoriae dicoriae, Phenacoccus, 49, 216, 235, 237, 238, 268, 496 didymus, Anisococcus, 3, 48, 55, 59, 61, 62, 63, 494 Dieldrin, 13 difficilis, Dysmicoccus, 157 Digger mealybug. See Scaptococcus californicus digitules, 26, 38 Dimefox, 13 Dimethoate (Cygon®), 14 diminutus, Trionymus, 4, 51, 461, 465, 466, 498 diodium, Dysmicoccus, 157 direct benefits (to mealybugs), 21 Discococcus, 18, 19, 25, 41, 42, 44, 46, 134, 136 caricis, 4, 49, 136, 137, 138, 140, 144, 495 flabellatus 4, 49, 136, 137, 139, 140, 495 graminis, 4, 44n, 49, 136, 140, 141, 142, 495 hopi, 4, 49, 136, 142, 143, 147, 495 simplex, 4, 49, 136, 144, 145, 495 spectabilis, 18, 19, 49, "136, 142, 144, 146, 147, 495 discoid (pores), 39 discoidal pores, 38, 40, 56 disjunctus, Rhizoecus, 46n, 50, 371, 372, 385, 386, 387, 497 Dispar mealybug. See Pseudococcus dispar dispar, Pseudococcus, 50, 289, 291, 292, 293, 326, 496 Distichlicoccus, 41, 45, 147, 148 arundinis, 49, 148, 149, 150, 152, 154, 495 californicus, 4, 49, 148, 150, 151, 152, 154, 495 megacirculus, 49, 148, 150, 152, 153, 154, 495 salinus, 4, 18, 49, 148, 152, 154, 155, 156, 495 distichlii, Tridiscus, 4, 51, 455, 456, 498 distichlium, Paludicoccus, 4, 49, 211, 212, 213, 495 distichlium, Sphaerococcus, 211 DISTRIBUTION TABLE, 4 9 3

Di-Syston®, 13 Diverse mealybug. See Pseudococcus diversus

diversus, Pseudococcus, 50, 288, 291, 293, 294, 295, 496 Dolichoderus, 22 Dolus mealybug. See Trionymus dolus dolus, Trionymus, 4, 17, 51, 461, 467, 468, 498 dorsal ostioles, 39 dorsal setae, 40 Dorthezia, 280 Douglas fir mealybug. See Puto profusus drenching, 14 dubia, Dactylopius, 208 ducts, 39, 40 Dusky grass mealybug. See Trionymus furvus Dysmicoccus, 40, 41, 47, 156, 162, 288 aciculus, 4, 49, 157, 158, 159, 495 brevipes, 10, 13, 24, 29, 49, 157, 160, 161, 162, 495 mackenziei, 49, 157, 162, 163, 164, 495 pinicolus, 18, 49, 157, 164, 165, 166, 495 quericolus, 4, 49, 158, 166, 167, 168, 495 racemus, 49, 157, 168, 169, 170, 495 ryani, 4, 49, 157, 168, 170, 171, 172, 495 timberlakei, 4, 49, 157, 172, 173, 174, 495 vacuatus, 49, 157, 174, 175, 176, 495 Echeveria mealybug. See Phenacoccus echeveriae echeveriae, Phenacoccus, 22, 49, 215, 238, 239, 240, 248, 496 echinatus, Puto, 8, 26, 50, 328, 347, 348, 349, 350, 497 ECOLOGY 16 ECONOMIC IMPORTANCE,

6

Ehrhornia, 41, 46, 134, 136, 140, 176 cupressi, 4, 18, 49, 176, 177, 178, 495 Ehrhornia group, 41 eluminatus, Rhizoecus, 51, 372, 387, 388, 389, 400, 497 Endrin, 13 Ephedra mealybug. See Anisococcus ephedrae ephedrae, Aniscoccus, 3, 48, 55, 63, 64, 65, 494 epiphyllis, Rhizoecus, 379 Eremicus mealybug. See Phenacoccus eremicus eremicus, Phenacoccus, 4, 17, 50, 214n, 215, 240, 241, 242, 248, 255, 496 Eriococcus, 52, 130 eriogoni, Phenacoccus, 4, 50, 214, 235, 242, 243, 244, 266, 496

GENERAL eriogoni, Spilococcus, 4, 16, 51, 411, 423, 424, 425, 498 Eriogonum mealybug. See Spilococcus eriogoni Erium, 52 Erium confusum, 52 erythrosin, 34, 35 eschscholtziae, Phenacoccus, 5, 17, 19, 50, 216, 244, 245, 246, 247, 496 Essig's aphid fluid, 34, 35 euchromatic, 30 Eurycoccus, 47, 81 eyes, 25, 38 falcifer, Rhizoecus, 4, 9, 10, 17, 29, 41, 43, 51, 370, 371, 389, 390, 391, 497 False obscure mealybug. See Pseudococcus pseudobscurus Farinococcus, 80, 81 farinosus, Dactylopius virgatus variety, 179, 181 Felt sac mealybug. See Cryptoripersia salina felted sac, 42 Ferns, 19 Ferris atriplex mealybug. See Humococcus atriplicis Ferris grass mealybug, See Discococcus graminis Ferris oak mealybug. See Dysmicoccus quericollus Ferrisia, 41, 42, 45, 179, 182 virgata, 4, 5, 7, 10, 13, 25, 42, 49, 179, 180, 181, 182, 495 Ferrisia group, 41 Ferrisiana, 179 festucae, Trionymus, 4, 17, 51, 461, 469, 470, 498 FIELD AND LABORATORY METHODS, 3 2 FIELD CHARACTERISTICS MEALYBUG GENERA,

OF 42

Field key to common California Mealybugs, 10 Fifth instar male (adult), 25 filamentation, 43 filamentosus, Nipaecoccus, 206 fimbriatulus, Dysmicoccus, 157 First instar nymph, 24 fistulosus, Chorizococcus, 48, 494 flabellatus, Discococcus, 4, 49, 136, 137, 139, 140, 495 flaveolus, Phenacoccus, 215 floridana, Ferrisia, 179 Fluffy mealybug. See Puto echinatus foliage, 32, 43 fontanus, Distichlicoccus, 148, 150 Formica sp., 132, 227, 230 Formica camponoticeps, 204

Formica pilicornis, 107, 109, 224, 225, 254 Formica subpolita, 23, 309, 310 Formica subpolita (fusca group), 227, 230 Formica subpolita camponoticeps 132, 227, 230 Fourth instar female, 24 Fourth instar male, 25 fragilis, Pseudococcus, 4, 5, 6, 10, 26, 50, 172, 289, 295, 296, 298, 496 franseriae, Phenacoccus, 215 Frontal grass mealybug. See Trionymus frontalis frontalis, Trionymus, 51, 461, 471, 472, 498 fumigant control, 13 fungus, 20, 21 furvus, Trionymus, 51, 460, 473, 474, 498 gahani, Pseudococcus, 295 genacerores (pores), 39 generations, 27 GENERA (North American Pseudococcidae), 41 genes, 29 GENETIC

SYSTEMS,

28

geniculate, 38 geniculatus, Rhizoecus, 385, 387 genitalia, 26 Geococcus, 25, 38, 42, 44 Geococcus coffeae, 44n geraniae, Spilococcus 51, 411, 427, 428, 498 Geranium mealybug. See Spilococcus geraniae Giant mealybug. See Phenacoccus giganteus giganteus, Phenacoccus, 5, 50, 215, 248, 249, 496 giganticoxa, Pseudantonina, 286 globosus, Rhizoecus, 368 Golden mealybug. See Nipaecoccus aurilanatus gossypii, Phenacoccus, 4, 5, 7, 10, 13, 24, 25, 26, 27, 45n, 50, 215, 248, 250, 251, 252, 255, 271, 496 gossypii, Phenacoccus helianthi variety, 248 gracilis, Rhizoecus, 16, 17, 51, 371, 392, 393, 497 graminicola, Heterococcus, 189, 191, 193 graminis, Antonina, 4, 7, 8, 10, 13, 18, 29, 48, 74, 76, 77, 80, 494 graminis, Discococcus, 4, 44n, 49, 136, 140, 141, 142, 495 graminis, Ehrhornia, 136 Graminis ground mealybug. See Rhizoecus graminis graminis, Rhizoecus, 4, 51, 372, 394, 395, 497

graminosus, Phenacoccus, 5, 8, 10, 18, 19, 50, 215, 252, 255, 256, 257, 496 Grape mealybug. See Pseudococcus maritimus Grasses, 17, 33 grass crown, 42 Grass puto mealybug. See Puto acirculus grass root, 33 grindeliae, Trionymus, 85 Ground mealybug. See Rhizoecus falcifer Ground mealybug control, 13, 14 guatemalensis, Cataenococcus, 81 gum-chloral hydrate, 34 Gutierrezia mealybug. See Spilococcus gutierreziae gutierreziae, Spilococcus, 4, 51, 412, 429, 430, 498 Haanchen barley mealybug. See Trionymus haancheni haancheni, Trionymus, 8, 10, 47n, 51, 461, 475, 476, 498 HABITATS (MEALYBUG),

16

halteres, 26 harbisoni, Phenacoccus, 214, 252 HCN, 14 Head, 36 helianthi, Penacoccus, 4, 50, 215, 224, 257, 258, 259, 496 Heliococcus, 41, 42, 44, 179, 181, 182, 186 adenostomae, 49, 183, 185, 495 atriplicis, 49, 182, 183, 185, 186, 188, 495 stachyos, 4, 19, 49, 183, 185, 186, 188, 495 Heliococcus group, 41 hermaphroditism, 27 heterochromatic, 30 heterochromatization, 29n Heterococcus, 41, 42, 44, 188, 189, 204 arenae, 4, 18, 49, 188, 189, 190, 191, 495 pulverarius, 4, 5, 8, 10, 13, 14, 42, 49, 189, 191, 192, 193, 495 Heterococcus group, 41 Hilaria mealybug. See Humococcus hilariae hilariae, Humococcus, 4, 49, 197, 198, 199, 495 hilariae, Ripersia, 193 hind coxa, 38 Hippeococcus, 22 hirsutus, Maconellicoccus, 13 HISTORY, 1

honeydew, 12, 21, 22, 25 hopi, Discococcus, 4, 49, 136, 142, 143, 147, 495 Hopi mealybug. See Discococcus hopi hosing the plants, 14

INDEX

HOST POSITIONS ( O F MEALYBUGS),

521

19

humilts, Dactylopius virgatus variety, 179, 181 Humococcus, 42, 44, 45, 47, 91, 148, 193, 203 atriplicis, 4, 7, 19, 49, 194, 195, 197, 199, 201, 495 caritus, 49, 194, 196, 197, 495 hilariae, 4, 49, 194, 197, 198, 199, 495 inornatus, 49, 194, 199, 200, 201, 495 hurdi, Dysmicoccus, 158 hurdi, Phenacoccus, 215 hymenocleae, Amonostherium, 52, 54 Hypogeic mealybugs, 9, 371 Hypogeococcus, 44, 46, 201 spinosus, 4, 49, 201, 202, 203, 495 hypolithus, Cryptoripersia, 130 Imperfect mealybug. See Phenacoccus defectus Imperial mealybug. See Anisococcus imperialis imperialis, Anisococcus, 48, 55, 65, 66, 67, 494 Implicated mealybug. See Spilococcus implicatus implicatus, Spilococcus, 4, 5, 18, 51, 170, 412, 431, 432, 433, 498 importatus, Pseudococcus, 50, 288, 298, 299, 300, 311, 313, 320, 496 Imported mealybug. See Pseudococcus importatus Inanimate objects, 19 incomptus, Phenacoccus, 216, 247 indecisus, Chorizococcus, 45, 89 indica, Antonina, 76 Infernal mealybug. See Phenacoccus infernalis infernalis, Phenacoccus, 50, 215, 219, 242, 255, 260, 261, 496 insignis, Heliococcus, 183 Instars and Generations, 27 Inornate mealybug. See Humococcus inornatus inornatus, Humococcus, 49, 194, 199, 200, 201, 495 Interior live oak mealybug. See Anisococcus quercus Interrupted mealybug. See Chorizococcus interrupts interruptus, Chorizococcus, 49, 89, 101, 102, 103, 105, 117, 494 INTRODUCTION, 1

invaginated (duct), 40 irishii, Chorizococcus, 89 Japanese mealybug. See Planococcus kraunhiae

522

GENERAL

junceus, Dysmicoccus, 157 juniperi, Spilococcus, 412 Keifer mealybug. See Spilococcus keiferi keiferi, Spilococcus, 51, 412, 434, 435, 498 Kellogg mealybug. See Radicoccus kelloggi kelloggi, Radicoccus, 4, 50, 368, 369, 370, 497 Kermicus, 76 KEYS

NORTH

AMERICAN

SPECIES, 4 8 KEY T O N O R T H

TO

AMERICAN

GENERA,

44

kingii, Pseudococcus, 289 Kondo mealybug. See Rhizoecus kondonis kondonis, Rhizoecus, 4, 5, 9, 13, 17, 51, 371, 394, 396 398, 497 kraunhiae, Planococcus, 4, 50, 280, 282, 283, 284, 285, 496 Kuwana brown mealybug. See Paludicoccus distichlium Kuwana pine mealybug. See Crisicoccus pini kuwanae, Phenacoccus, 268 labium, 38 Large-celled mealybug. See Anisococcus didymus Large-circulus mealybug. See Distichlicoccus megacirculus Large-duct mealybug. See Phenacoccus megaulus Large-setae mealybug. See Pseudococcus megasetosus Large yucca mealybug. See Puto yuccae larreae, Spilococcus 4, 19, 51, 411, 436, 437, 498 lasii, Dysmicoccus, 157 lasiorum, Puto, 329 Lasius americanus, 398 Lasius flavus, 21, 230 laticribellum, Puto, 18, 26, 50, 328, 343, 350, 351, 352, 497 latipes, Pseudococcus longisptnus variety, 313, 316 Lead arsenate, 14 "lecanoid" chromosome system, 28, 29 ledi, Nipaecoccus, 206 legs, 25, 26, 38, 39, 40 Lenania, 25 Leptomastidea abnormis, 6 leucosomus, Rhizoecus, 4, 51, 371, 398, 399, 400, 402, 497 lichtensioides, Amonostherium, 3, 16, 48, 52, 53, 54, 494 lichtensioides, Dactylopius, 52 L I F E HISTORY, 2 4

lignin pink, 34, 35

INDEX Lily bulb mealybug. See Chorizococcus lounsburyi Lindane, 13 LIST O F CALIFORNIA M E A L Y BUGS, 4 8 LIST OF COLORED PHOTOGRAPHS, 4 LITERATURE CITED, 499

loculi, 38 Long-setae mealybug. See Pseudococcus longisetosus Long-tailed mealybug. See Pseudococcus longispinus longirostralis, Pseudococcus, 160 longifilis, Dactylopius, 303 longisetosus, Pseudococcus, 4, 50, 289, 300, 301, 302, 309, 496 longispinus, Pseudococcus, 4, 7, 10, 13, 16, 19, 26, 29, 50, 288, 303, 304, 302, 309, 496 lotearum, Phenacoccus, 50, 216, 262, 263, 496 Lotus mealybug. See Phenacoccus lotearum lounsburyi, Chorizococcus, 5, 49, 89, 101, 103, 104, 105, 494 lounsburyi, Trionymus, 105 lycii, Phenacoccus, 4, 50, 214, 235, 264, 265, 496 mackenziei, Dysmicoccus 49, 157, 162, 163, 164, 495 Macrocerococcus, 326 MACRO-ENVIRONMENTAL PREFERENCES, 16

Maconellicoccus hirsutus, 13 macropterous, 26 macswaini, Pseudococcus, 289, 293, 295 magnus, Trionymus, 47n, 461 malacearum, Pseudococcus, 313, 316 Malathion, 12, 14 malvastrus, Heliococcus, 181 Manica bradleyi, 130 Many-pored mealybug. See Chorizococcus polyporus Manzanita mealybug. See Puto arctostaphyli marchali, Pseudococcus, 179 maritimus, Pseudococcus, 5, 6, 7, 10, 12, 13, 50, 289, 306, 307, 308, 309, 316, 496 maritimus-malacearum (complex), Pseudococcus, 308, 315 maritimus, Rhizoecus, 371 mayanus, Rhizoecus, 372 mcdanieli, Dysmicoccus, 157 McKenzie adenostoma mealybug. See

Heliococcus adenostomae McKenzie mealybug. See Dysmicoccus mackenziei McKenzie pine mealybug. See Dysmicoccus pinicolus McKenzie salt grass mealybug. See Humococcus caritus megacirculus, Distichlicoccus, 49, 148, 150, 152, 153, 154, 495 megasetosus, Pseudococcus, 50, 288, 309, 310, 311, 496 megaulus, Phenacoccus, 50, 214, 244, 266, 267, 496 menkei, Rhizoecus, 371 merrilli, Dysmicoccus, 157 Mesquite mealybug. See Spilococcus prosopidis Mexican mealybug. See Phenacoccus gossypii mexicanus, Puto, 329 microcirculus, Pseudococcus, 7, 10, 50, 288, 300, 311, 312, 313, 496 MICRO-ENVIRONMENTAL PREFERENCES, 16

microporus, Chorizococcus, 103, 105 migration, 20 milleri, Phenacoccus, 214, 244, 247 Miller sand dune mealybug. See Scaptococcus milleri milleri, Scaptococcus, 5, 19, 51,407, 498 Mimic mealybug. See Puto mimicus mimicus, Puto, 50, 327, 352, 353, 354, 497 Minimus mealybug. See Phenacoccus minimus minimus, Phenacoccus, 4, 50, 216, 238, 240, 242, 268, 269, 496 minor, Pseudococcus calceolaria variety, 280 minute circular (pores), 39, 56n miscellaneous chemical control, 14 Misericoccus, 18, 25, 42, 44, 203 arenarius, 4, 19, 42, 49, 204, 205, 495 Mocker grass mealybug. See Trionymus mocus mocus, Trionymus, 4, 51, 461, 477, 478, 498 Modoc grass mealybug. See Trionymus modocensis modocensis, Trionymus, 4, 47n, 51, 461, 479, 480, 498 Monterey pine mealybug. See Dysmicoccus aciculus morphological structures 36, 37, 38-40

MORPHOLOGY, 3 6

Morrison pygmy mealybug. See Pygmaeococcus morrisoni Morrisonella, 370, 372, 383, 394 morrisoni, Dysmicoccus, 158 morrisoni, Pygmaeococcus, 50, 366, 367, 368, 497 moruliferus, Rhizoecus, 389 mouthparts, 38 muhlenbergiae, Humococcus, 194, 197 multiocular disk (pores), 25, 39 MULTIPLE

HOST

POSITIONS,

19 multiorbis, Tridiscus, 4, 51, 457, 458, 498 Multiple circulus mealybug. See Tridiscus multiorbis Multiple-setae mealybug. See Phenacoccus multisetosus multisetosus, Phenacoccus, 50, 215, 252, 268, 270, 271, 496 Myers mealybug. See Trionymus myersi myersi, Trionymus, 51, 460, 481, 482, 498 Myrmecocystus semirufus 253 Naiacoccus, 52 nanus, Trionymus, 460 nemoralis, Rhizoecus, 372, 385, 400 neobrevipes, Dysmicoccus, 157, 160, 164 neomexicanus, Chorizococcus, 88 neomexicanus, Rhizoecus, 372, 394 Neorhizoecus, 42, 46 Nest mealybug. See Anisococcus oregonensis New Zealand Flax mealybug. See Trionymus diminutus Nicotine, 14 nigeriensis, Heterococcus, 14 nipae, Dactylopius, 206 nipae, Nipaecoccus, 4, 5, 49, 206, 208, 209, 210, 211 495 Nipaecoccus, 46, 52, 201, 206 aurilanatus, 4, 5, 49, 206, 207, 208, 210, 495 nipae, 4, 5, 49, 206, 208, 209, 210, 211, 495 njalensis, Planococcoides, 10, 13 nonarius, Phenacoccus, 214, 216 nonbiological objects, 19 NORTH

AMERICAN

41-42 Key to, 44

GENERA,

GENERAL North coast mealybug. See Trionymus festucae nortoni, Antonina, 74 Notable mealybug. See Discococcus spectabilis Noxious bamboo mealybug. See Antonina pretiosa nuclei, 29 nudus, Phenacoccus, 191 Nulliporus mealybug. See Puto nulliporus nulliporus, Puto, 50, 327, 352 354, 355, 357, 497 obesus, Dysmicoccus, 13, 157 Obscure mealybug. See Pseudococcus obscurus occidentals, Heterococcus, 189 Oecophylla, 21 oil emulsions, 14 olivaceous, Cataenococcus, 4, 5, 48, 81, 82, 83, 494 olivaceus Dactylopius, 80 Olive-brown mealybug. See Cataenococcus olivaceus omniverae, Pseudococcus, 306, 308 oogenesis, 31 Oracella, 47 oral-collar (duct), 40 oral-rim (duct), 40 Orchid mealybug. See Pseudococcus microcirculus oregonensis, Anisococcus, 4, 5, 48, 55, 67, 68, 70, 494 Organic phosphates, 12 osborni, Heliococcus, 183, 188 ostioles, 21, 25, 40 oviparous, 42 Oviposition, 24 ovisac, 24, 42, 43 ovoviviparous, 24, 42 Pacific mealybug. See Puto pacificus pacificus, Puto, 17, 26, 50, 328, 354 356, 359, 497 palmarum, Palmicola, 26 Palmicola palmarum, 26 Paludicoccus, 41, 44, 211 distichlium, 4, 49, 211, 212, 213, 495 Paradoxococcus, 41, 47 Parker mealybug. See Spilococcus, parkeri Parathion, 12, 14 parkeri, Spilococcus, 51, 411, 438, 439, 498 Paroudablts, 213, 257 Parthenogenesis, 27, 29, 30, 31 parvicirculus, Spilococcus, 51, 411, 440, 441, 498 patulae, Dysmicoccus 158 Pauper mealybug. See Phenacoccus pauperatus pauperatus, Phenacoccus, 4, 50, 216, 234, 262, 271, 272, 273, 496

Pear-shaped mealybug. See Misericoccus arenarius Pedronia, 19 Peliococcus, 41, 45 Peliococcus group, 41 Penultimate mealybug. See Chorizococcus penultimatus penultimatus, Chorizococcus, 49, 89, 105, 106, 107, 494 peregrinus, Chorizococcus, 103, 105 peregrinus, Pseudococcus (Trionymus), 103 Perennial-flowering plants (excluding grasses), 16 Perforated mealybug. See Pseudococcus pertusus Pergandiella, 459, 460, 461 Pergrolhula, 459 Permanent mounts, 34 perrisii, Westwoodia, 460 pertusus, Pseudococcus, 50, 288, 318, 319, 320, 496 Peukinococcus, 213 Phaenococcus, 213 phenacocciformis, Pseudococcus citri variety, 280 Phenacoccus, 16, 25, 41, 43, 44, 45, 46, 179, 182, 186, 213, 214, 257, 363 alleni, 16, 21, 49, 215, 216, 217, 218, 219, 240, 242, 260, 495 artemisiae, 4, 16, 17, 20, 49, 214, 219, 220, 221, 495 cajonensis, 49, 214, 221, 222, 223, 225, 259, 495 celtisifoliae, 4, 5, 49, 216, 225, 226, 495 colemani, 4, 5, 19, 23, 49, 216, 225, 227, 228, 229, 230, 234, 248 (misident.), 496 dearnessi, 4, 49, 214, 231, 232, 496 defectus, 4, 16, 49, 215, 231, 233, 234, 275, 279, 496 destitutus, 49, 214, 235, 236, 264, 496 dicoriae, 49, 216, 235, 237, 238, 268, 496 echeveriae, 22, 49, 215, 238, 239, 240, 248, 496 eremicus, 4, 17, 50, 214n, 215, 240, 241, 242, 248, 255, 496 eriogoni, 4, 50, 214, 235, 242, 243, 244, 266, 496 eschscholtziae 5, 17, 19, 50, 216, 244, 245, 246, 247, 496 giganteus, 5, 50, 215, 248, 249, 496 gossypii, 4, 5, 7, 10, 13, 24, 25, 26, 27 45n, 50, 215, 248, 250, 251, 252, 271, 496 graminosus, 5, 8, 10, 18, 19, 50, 215, 252, 255, 256, 257, 496

helianthi, 4, 50, 215, 224, 257, 258, 259, 496 infernalis, 50, 215, 219, 242, 255, 260, 261, 496 lotearum, 50, 216, 262, 263, 496 lycii, 4, 50, 214, 235,264, 265, 496 megaulus, 50, 214, 244, 266, 267, 496 minimus, 4, 50, 216, 238, 240, 242, 268, 269, 496 multisetosus, 50, 215, 252, 268, 270, 271, 496 pauperatus, 4, 50, 216, 234, 262, 271, 272, 273, 496 solani, 4, 5, 17, 19, 50, 216, 230, 234, 273, 274, 277, 279, 496 solenopsis, 4, 19, 50, 215, 275, 277, 278, 279, 496 Phenacoccus group, 41 phoradendri, Cataenococcus, 81 Phorate (Thimet®), 13 phyllococcus, Lecanium, 280 phylogenetic, 41 picrospinus, Phenacoccus, 225 pigmentation, 39 Pine bark mealybug. See Puto laticribellum Pineapple mealybug. See Dysmicoccus brevipes pini, Crisicoccus, 4, 18, 49, 125, 127, 128, 129, 494 pini, Dactylopius, 125 pinicolus, Dysmicoccus, 18, 49, 157, 164, 165, 166, 495 Planococcoides njalensis, 10, 13 Planococcus, 41, 43, 46, 280 citri, 4, 5, 6, 7, 10 13, 14, 16, 24, 26, 27, 29, 43, 50, 280, 281, 282, 283, 285, 496 kraunhiae, 4, 50, 280, 282, 283, 284, 285, 496 Planococcus group, 41 plantar surface, 38 plowing, 14 polyploid cells, 31 polyporus, Chorizococcus, 17, 19, 49, 89, 96, 107, 108, 109, 494 polyvinyl alcohol, 34 porcelain casserole, 34 pore types, 39 pores, 39 Porococcus, 41, 46 Porous mealybug. See Chorizococcus fistulosus porterae, Humococcus, 193 PREPARATION OF SLIDE MOUNTS,

34

Preserve plant material, 33 Pressus mealybug. See Spilococcus pressus pressus, Spilococcus, 51, 412, 442, 443, 498 pretiosa, Antonina, 4, 48, 74, 76, 78, 79, 80, 494

INDEX

523

Price mealybug. See Puto pricei pricei Puto, 50, 328, 337, 357, 358, 359, 497 primary oocyte, 31 Pritchard mealybug. See Rhizoecus pritchardi pritchardi, Rhizoecus, 5, 9, 10, 13, 14, 19, 51, 372, 387, 400, 401, 402, 497 probrevipes, Dysmicoccus, 157 profusus, Puto, 50, 328, 360, 361, 497 prosopidis, Spilococcus, 4, 51, 412, 444, 445, 498 protection from enemies, 20 prunicolus, Pseudococcus, 289 Pseudantonina, 41, 45, 285, 286 arundinariae, 4, 50, 286, 287, 496 pseudobrevipes, Dysmicoccus, 160 pseudobrevipes, Pseudococcus, 160. . pseudobscurus, Pseudococcus, 50, 289, 320, 321, 322, 496 PSEUDOCOCCIDAE (Definition), 40 PSEUDOCOCCIDAE (North American Genera), 41 Pseudococcus, 52, 56, 59, 63, 67, 81, 85, 103, 125, 127, 154, 156, 160, 166, 170, 172, 179, 206, 208, 210, 288, 343, 465 aberrans, 50, 288, 289, 290, 291, 496 dispar, 50, 289, 291, 292, 293, 326, 496 diversus, 50, 288, 291, 293, 294, 295, 496 fragilis, 4, 5, 6, 10, 26, 50, 172, 289, 295, 296, 298, 496 importatus, 50, 288, 298, 299, 300, 311, 313, 320, 496 longisetosus, 4, 50, 289, 300, 301, 302, 309, 496 longispinus, 4, 7, 10, 13, 16, 19, 26, 29, 50, 288, 303, 304, 305, 496 maritimus, 5, 6, 7, 10, 12, 13, 50, 289, 306, 307, 308, 309, 316, 496 megasetosus, 50, 288, 309, 310, 311, 496 microcirculus. 7. 10. 50. 288, 300, 311, 312, 313, 496 obscurus 5, 6, 16, 29, 50, 288, 302, 306 (misident.), 308, 313, 314, 315, 316, 318, 322, 496 pertusus, 50, 288, 318, 319, 320, 496 pseudobscurus, 50, 289, 320, 321, 322, 496 sorghiellus, 50, 288, 322, 323, 324, 496

524

GENERAL

Pseudococcus (Cont.) sparsus, 50, 289, 293, 324, 325, 326, 497 Pseudococcus group, 41 Pseudolecanium, 211 pseudonipae, Dactylopius, 208, 210 pseudonipae, Pseudococcus, 208 psoraleae, Chorizococcus, 49, 89, 91, 101, 109, 110, 111, 494 pulverarius, Heterococcus, 4, 5, 8, 10, 13, 14, 42, 49, 189, 191, 192, 193, 495 Purple mealybug. See. Chorizococcus californicus purpurea, Antonina, 73 Puto, 16, 18, 25, 26, 27, 28, 29, 41, 42, 43, 45, 59, 214, 326, 327, 329, 350 acirculus, 5, 50, 327, 329, 330, 332, 497 albicans, 8, 50, 329, 331, 332, 333, 497 ambiguus, 4, 26, 50, 327, 328, 333, 334, 335, 339, 345, 359, 497 arctostaphyli, 4, 26, 50, 329, 332, 335, 336, 337, 497 atriplicis, 50, 328, 337, 338, 339, 497 bryanthi, 4, 50, 328, 335, 339, 342, 359, 497 californicus, 5, 50, 328, 339, 341, 342, 497 cupressi, 4, 18, 50, 328, 343, 344, 345, 350, 357, 360, 497 decorosus, 17, 26, 50, 328, 345, 346, 497 echinatus, 8, 26, 50, 328, 347, 348, 349, 350, 497 laticribellum, 18, 26, 50, 328, 343, 350, 351, 352, 497 mimicus, 50, 327, 352, 353, 354, 497 nulliporus, 50, 327, 352, 354, 355, 357, 497 pacificus, 17, 26, 50, 328, 354, 356, 359, 497 pricei, 50, 328, 337, 357, 358, 359, 497 profusus, 50, 328, 360, 361, 497 simmondsiae, 50, 328, 347, 360, 362, 363, 365, 497 yuccae, 4, 17, 19, 26, 50, 59, 329, 333 (misident.), 335, 347, 350, 363, 364, 365, 497 Puto group, 42 Putonia, 326 (preoccupied) Pygmaeococcus, 25, 38, 40, 41, 42, 46, 366 morrisoni, 50, 366, 367, 368, 497

INDEX quadricirculus, Trionymus, 47n, 51, 461, 483, 484, 498 quaintancei, Ferrisia, 179 quercicolus, Dysmicoccus, 4, 49, 158, 166, 167, 168, 495 quercinus, Spilococcus, 51, 411, 446, 447, 498 quercus, Anisococcus, 4, 5, 48, 55, 56, 67, 69, 70, 71; 494 quercus, Pseudococcus, 166 (misident.) quinquelocular pores, 25, 39 racemus, Dysmicoccus, 49, 157, 168, 169, 170, 495 Radicoccus, 42, 46, 368, 370 kelloggi, 4, 50, 368, 369, 370, 497 ramonae, Phenacoccus, 363 rectal cavity, 21 reducta, Chorizococcus, 4, 49, 88, 111, 112, 113, 123. 494 Redwood mealybug. See Spilococcus sequoiae regnillhoi, Phenacoccus, 231 resistance, 12 Rhizoecus, 9, 10, 16, 18, 25, 33, 38, 41, 42, 43, 46, 366, 370, 371, 376, 383, 387, 391 associatus, 50, 372, 373, 374, 497 bicirculus, 50, 371, 374, 375, 497 bituberculatus, 50, 371, 376, 377, 404, 497 browni, 50, 371, 376, 378, 379, 497 cacticans, 4, 5, 50, 371, 374, 379, 380, 381, 497 californicus, 4, 50, 371, 381, 382, 383, 497 cyperalis, 4, 50, 372, 383, 384, 385, 400, 497 disjunctus, 46n, 50, 371, 372, 385, 386, 387, 497 eluminatus, 51, 372, 387, 388, 389, 400, 497 falcifer, 4, 9, 10, 17, 29, 41, 43, 51, 370, 371, 389, 390, 391, 497 gracilis, 16, 17, 51, 371, 392, 393, 497 graminis, 4, 51, 372, 394, 395, 497 kondonis, 4, 5, 9, 13, 17, 51, 371, 394, 396, 398, 497 leucosomus, 4, 51, 371, 398, 399, 400, 402, 497 pritchardi, 5, 9, 10, 13, 14, 19, 51, 372, 387, 400, 401, 402, 497 sonomae, 51, 371, 383, 398, 402, 403, 404, 497 spinosus, 51, 371, 376, 397, 404, 405, 406, 497 Rhizoecus group, 42 Rhodania, 129 Rhodesgrass mealybug. See Antonina graminis

Ripersia, 86, 191, 197, 204, 469 Ripersia confusella, 83 (misident.) Ripersia hilariae, 193 Ripersia trichura, 129 Ripersia villosa, 83, 86 Ripersiella, 368, 379, 398 root infestations, 17 roots, 33, 42 roseotinctus, Dysmicoccus, 157, 174 rostellum, Chorizococcus, 4, 18, 19, 49, 88, 113, 114, 115, 494 rostellum, Trionymus, 29 rostrum, 38 Rotenone, 14 rubivorus, Phenacoccus, 215 ryani, Dysmicoccus, 4, 49, 157, 168, 170, 171, 172, 495 ryania, 14 Ryegrass mealybug. See Phenacoccus graminosus sacchari, Saccharicoccus, 26, 27, 29 Saccharicoccus, 46 Sagebrush mealybug. See Amonostherium lichtensioides salina, Cryptoripersia, 4, 5, 22, 42, 49, 130, 131, 132, 494 salinus, Dactylopius, 147 salinus, Distichlicoccus, 4, 18, 49, 148, 152, 154, 155, 156, 495 salmonaceus, Dysmicoccus, 158 Salt grass mealybug. See Distichlicoccus salinus Saltbrush mealybug. See Heliococcus atriplicis Sand mealybug. See Heterococcus arenae sandini, Puto, 18, 24, 27, 328, 342 Scaptococcus, 41, 43, 46, 52, 406 californicus, 19, 51, 407, 497 milleri, 5, 19, 51, 407,498 Schradan ( O M P A ) , 13 SCOPE OF STUDY, 4

Second instar nymph, 24 secondary oocyte, 31 Sedge mealybug. See Discococcus caricis Segmentation, 25, 36 senarius, Chorizococcus, 49, 89, 115, 116, 117, 494 sequoiae, Pseudococcus, 343 sequoiae, Spilococcus, 4, 51, 412, 448, 449, 498 serrata, Ripersia, 208 setosa, Ferrisia, 179 sex determination, 29 sex ratio, 27, 28 shaferi, Chorizococcus, 89 Short-antennae mealybug. See Chorizococcus senarius

Short-legged mealybug. See Chorizococcus brevicruris Signoretia, 459 (preoccupied) Simonds mealybug. See Anthelococcus simondsi simondsi, Anthelococcus, 48, 71, 72, 73, 494 Simmondsia mealybug. See Puto simmondsiae simmondsiae, Puto 50, 328, 347, 360, 362, 363, 365, 497 simple disc pores, 39 Simple mealybug. See Discococcus simplex simplex, Discococcus, 4, 49, 136, 144, 145, 495 simplex, Phenacoccus, 363 Six-circuli mealybug. See Discococcus flabellatus Slender ground mealybug. See Rhizoecus gracilis slide mounts, 34 Small circulus mealybug. See Spilococcus parvicirculus Small mealybug. See Dysmicoccus vacuatus Small yucca mealybug. See Chorizococcus yuccae Smith grass mealybug. See Trionymus smithii smithii, Trionymus, 4, 17, 51, 105, 107, 460, 485, 486, 498 smutting, 12 Snelling mealybug. See Chorizococcus snellingi snellingi, Chorizococcus, 49, 88, 117, 118, 119, 494 solani, Phenacoccus, 4, 5, 17, 19, 50, 216, 230, 234, 273, 274, 277, 279, 496 solani, Pseudococcus, TTi (misident.) solani, Rhizoecus, 371, 392 Solanum mealybug. See Phenacoccus solani Solenopsis mealybug. See Phenacoccus solenopsis Solenopsis geminata, 277 Solenopsis molesta validiuscula, 168, 277 Solenopsis truncorum, 262 solenopsis, Phenacoccus, 4, 19, 50, 215, 275, 277, 278, 279, 496 Sonoma ground mealybug. See Rhizoecus sonomae sonomae, Rhizoecus, 51, 371, 383, 398, 402, 403, 404, 497 sooty mold, 12, 21 sorghiellus, Pseudococcus, 50, 288, 322, 323, 324, 496 Southern California mealybug. See Spilococcus atriplicis Sparse mealybug. See Pseudococcus sparsus

GENERAL sparsus, Pseudococcus, 50, 289, 293, 324, 325, 326, 497 species replacement, 27 spectabilis, Discococcus, 18, 19, 49, 136, 142, 144, 146, 147, 495 spermatogenesis, 30 Sphaerococcus, 76, 176, 211 Spilococcus, 16, 25, 38, 41, 42, 45, 88, 111, 214, 343, 410 andersoni, 4, 51, 170, 412, 413, 498 atriplicis, 4, 17, 51, 412, 414, 415, 498 cactearum, 5, 24, 51, 411, 416, 417, 498 ceanothi, 51, 412, 419, 420, 498 corticosus, 16, 51, 411, 421, 422, 498 eriogoni, 4, 16, 51, 411, 423, 424, 425, 498 geraniae, 51, 411, 427, 428, 498 gutierreziae, 4, 51, 412, 429, 430, 498 implicatus, 4, 5, 18, 51, 170, 412, 431, 432, 433, 498 keiferi, 51, 412, 434, 435, 498 larrea, 4, 19, 51, 411, 436, 437, 498 parkeri, 51, 411, 438, 439, 498 parvicirculus, 51, 411, 440, 441, 498 pressus, 51, 412, 442, 443, 498 prosopidis, 4, 51, 412, 444, 445, 498 quercinus, 51, 411, 446, 447, 498 sequoiae, 4, 51, 412, 448, 449, 498 ventralis, 51, 411, 450, 451, 498 Spinose ground mealybug. See Rhizoecus spinosus Spinose mealybug. See Hypogeococcus spinosus spinosus, Hypogeococcus, 4, 49, 201, 202, 203, 495 spinosus, Rhizoecus, 51, 371, 376, 397, 404, 405, 406, 497 spiracles, 26, 38 Square circulus mealybug. See Trionymus quadricirculus Squawcarpet mealybug. See Spilococcus ceanothi stachyos, Heliococcus, 4, 19, 49, 183, 186, 187, 188, 495 staining solutions, 35 stangei, Rhizoecus, 372

steelii, Spilococcus, 411 stellate pores, 26 stems, 32 Stemmatomerinx, 41, 44, 453 decorata, 4, 51, 453, 454, 498 Strange aspects of antmealybug associations, 22 straussiae, Pseudococcus, 29 Striped mealybug. See Ferrisia virgata subterranean, 33, 42, 43 subterranean travel, 19 succession (seasonal), 27 Succession in mealybugs, 27 Sunflower mealybug. See Phenacoccus helianthi symbionts, 30 Syrmococcus, 18, 41, 46 Systemic insecticides, 13 Systox, 13 talini, Dactylopius, 179 Tapinoma sessile, 275 TAXONOMY O F SPECIES, 4 8

Temporary mounts, 34 tenuipes, Ripersia, 83 TEPP, 13 terrestris, Ripersia, 389 Tetracnemus peregrinus, 7 Tetracnemus pretiosus, 6 tetrahydrofuran, 34 texana, Pseudantonina, 286 texensis, Dysmicoccus, 158 thelytoky, 28, 29 Third instar nymph, 24 Thorax, 36, 38 tibiaegracilis, Phenacoccus, 227 Timberlake mealybug. See Dysmicoccus timberlakei timberlakei, Dysmicoccus, 4, 49, 157, 172, 173, 174, 495 tooth, 38 totonicopanus, Rhizoecus, 371, 392 townsendi, Spilococcus, 412 Toxaphene, 13 toxins, 12 Trabutina, 52 translucent dots (pores), 56 Trechocorys, 208 tribulus, Clavicoccus, 29 trichura, Cryptoripersia, 130 trichura, Ripersia, 129 Tridiscus 47, 455 distichlii, 4, 51, 455, 456, 498 multiorbis, 4, 51, 457, 458, 498 trifolii Chnaurococcus 5, 17, 19, 42, 48, 83, 84, 85, 86, 494 trilocular pores, 25, 39, 40

Trinity ground mealybug. See Rhizoecus bicirculus Trionymus, 16, 17, 41, 42, 44, 47, 71, 73, 85, 86, 103, 113, 150, 459 americanus, 4, 51, 461, 462, 498 caricis, 4, 51, 460, 463, 464, 498 diminutus, 4, 51, 461, 465, 466, 498 dolus, 4, 17, 51, 461, 467, 468, 498 festucae, 4, 17, 51, 461, 469, 470, 498 frontalis, 51, 461, 471, 472, 498 furvus, 51, 460, 473, 474, 498 haancheni, 8, 10, 47n, 51, 461, 475, 476, 498 mocus, 4, 51, 461, 477, 478, 498 modocensis, 4, 47n, 51, 461, 479, 480, 498 myersi, 51, 460, 481, 482, 498 quadricirculus, 47n, 51, 461, 483, 484, 498 smithii, 4, 17, 51, 105, 107, 460, 485, 486, 498 utahensis, 51, 460, 487, 488, 498 winnemucae, 51, 460, 490, 491, 498 tritubular (pore), 39 trivittata, Ripersia, 83 trivittatus, Chnaurococcus, 85 Trochanter mealybug. See Pseudococcus sorghiellus Truncate setae mealybug. See Stemmatomerinx decorata tubular ducts, 40 tubulata, Cryptoripersia, 45n, 49, 130, 132, 133, 494 Tubulate mealybug. See Cryptoripersia tubulata twigs, 43 two-circuli mealybug. See Phenacoccus dearnessi utahensis, Pseudococcus neomexicanus variety, 487 ulter, Puto, 328 Unipored mealybug. See Rhizoecus disjunctus Utah grass mealybug. See Trionymus utahensis utahensis, Trionymus, 51, 460, 487, 488, 498 vaccinii, Cucullococcus, 4, 49, 134, 135, 136, 494

INDEX

525

vacuatus, Dysmicoccus, 49, 157, 174, 175, 176, 495 vallis, Trionymus, 113 variabilis, Chorizococcus, 89, 107, 121 variabilis, Heterococcus, 191, 193 vastator, Nipaecoccus, 206 VC-13®, 14 ventralis, Spilococcus, 51, 411, 450, 451, 498 villosa, Chnaurococcus, 4, 42, 48, 83, 85, 86, 87, 494 villosa, Ripersia, 83, 86 violascens, Trionymus, 461 virgata, Ferrisia, 4, 5, 7, 10, 13, 25, 42, 49, 179, 180, 181, 182, 495 virgata, Ferrisiana, 179 virgatus Dactylopius, 179 viridula, Spilococcus, 412 Viviparous ground mealybug. See Rhizoecus cyperalis vulva, 25, 36, 39 Westwoodia, 459 (preoccupied), 460 White ground mealybug. See Rhizoecus leucosomus White mealybug. See Puto albicans White sage mealybug. See Anisococcus crawii Wilkey mealybug. See Chorizococcus wilkeyi wilkeyi, Chorizococcus, 49, 86, 119, 120, 121, 494 wilmattae, Phenacoccus, 216 Wilson mealybug. See Chorizococcus wilsoni wilsoni, Chorizococcus, 49, 88, 99, 101, 121, 123, 494 wings, 26 Winnemuca grass mealybug. See Trionymus winnemucae winnemucae, Trionymus, 51, 460, 490, 491, 498 Woolly oak mealybug. See Chnaurococcus villosa xylene, 35 Yellow mealybug. See Distichlicoccus californicus yuccae, Chorizococcus, 49, 88, 123, 124, 125, 494 yuccae, Puto, 4, 17, 19, 26, 50, 59, 329, 333 (misident.), 335, 347, 350, 363, 364, 365, 497 zygote, 31

ADDENDUM Pseudococcus comstocki (Kuwana), Comstock mealybug, Found established in California for first time. On August 23, 1967 heavy infestations of the Comstock mealybug, Pseudococcus comstocki (Kuwana) were found on fruitless mulberry, Morus alba (Moraceae), growing on a residential property at Porterville, Tulare County, California. The mealybug was originally described on mulberry in Japan in 1902. In 1917 it was observed in North America on mulberry and apple in New York. During the period from 1925-1944 the Comstock mealybug was reported as a serious pest to apple, pear, peach, and umbrella catalpa in several eastern states including Connecticut, Delaware, Georgia, New Jersey, New York, Ohio, South Carolina, Virginia, and West Virginia (Hough, 1925; Haeussler and Clancy, 1944; and Clancy, 1944). One published account of its economic importance (Cutright, 1939) indicated that an apple orchard of about 60 acres in southern Ohio was found to be infested, and that 80 large Grimes trees suffered almost a total loss of crop as far as commercial pack was concerned. At that time no effective insecticidal control for the Comstock mealybug was known. As a consequence biological control measures were undertaken by the United States Department of Agriculture, Agriculture Research Service, Entomology Research Division. Certain parasites imported from Japan during 1939-1941 were extremely effective in controlling this mealybug (Haeussler and Clancy, 1944; and Clancy, 1944). Extensive survey of the Porterville area in California is currently being conducted by the California State Department of Agriculture, Bureau of Entomology, and the Tulare County Department of Agriculture to delimit the infestation. In addition, the United States Department of Agriculture at Riverside plans to introduce into California those parasites which proved so effective in controlling this mealybug in the eastern United States. It is ultimately hoped that this pest may be eradicated or nullified by biological control methods from California. REFERENCES:

Clancy, D. W. 1944. Biology of Allotropa burrelli, a gregarious parasite of Pseudococcus comstocki. Jour. Agric. Res. 69 ( 4 ) : 159-167, illus. Cutright, C. R. 1939. Comstock's mealybug, Pseudococcus comstocki (Kuw.) on apple in Ohio. Jour. Econ. Ent. 32 (6):p. 888. Haeussler, G. J., and D. W. Clancy 1944. Natural enemies of Comstock mealybug in the Eastern States. Jour. Econ. Ent. 37 (4):503-509. Hough, H. S. 1925. Biology and control of Comstock's mealybug on the umbrella catalpa. Virg. Agric. Expt. Sta., Tech. Bull. 29:5-27, illus.