Monograph on Brachystelma and Ceropegia in India 9780429061233, 0429061234, 9780429588303, 0429588305, 9780429590245, 0429590245, 9780429592188, 0429592183, 9780367183813

Table of contents :
Content: Cover
Half Title
Title Page
Copyright Page
Table of Contents
Preface
Authors
1: Introduction
1.1 Introduction
References
2: Brachystelma Taxonomy
2.1 Introduction
2.2 Systematic Enumeration
References
3: Traditional Uses, Pharmacognostic, Phytochemical, Pharmacological, and In Vitro Propagation Studies in Brachystelma Species
3.1 Introduction
3.2 Edible Uses
3.3 Medicinal Uses
3.4 Pharmacognostical Studies
3.5 Phytochemical Studies
3.5.1 Antioxidant Activity
3.5.2 Nutritional and Mineral Studies
3.5.3 Bromatological Analysis
3.6 In Vitro Propagation Studies 3.6.1 In Vitro Propagation Studies of Brachystelma glabrum3.6.2 In Vitro Propagation Studies of B. ngomense, B. pulchellum, and B. pygmaeum
3.7 Conclusion
References
4: Ceropegia Taxonomy
4.1 Introduction
4.1.1 Botanical History of Indian Ceropegia
4.2 Systematic Enumeration
References
5: Traditional Uses, Phytochemistry, and Pharmacology of Ceropegia Species
5.1 Introduction
5.2 Ornamental Uses
5.3 Traditional Uses
5.3.1 Edible Uses
5.3.2 Traditional Medicinal Uses
5.4 Pharmacognostic Studies of Ceropegia Species
5.4.1 Powder Characteristics
5.4.1.1 Physicochemical Analysis 5.5 Phytochemistry of Ceropegia Species5.5.1 Chemical Constituents in Ceropegia
5.5.2 Major Bioactive Constituents of Ceropegia Species
5.5.3 Chemical Synthesis of Cerpegin
5.6 Pharmacology of Ceropegia Species
5.6.1 Antimicrobial Activity
5.6.2 Antiurolithic Activity
5.6.3 Analgesic Activity
5.6.4 Hepatoprotective and Antioxidant Activity
5.6.5 Antioxidant Activity
5.6.6 Anticancer Activity
5.6.7 Antidiabetic Activity
5.6.8 Anti-inflammatory Activity
5.6.9 Gastroprotective Effect
5.6.10 Inhibitory Activity against Peptidases
References 6: Propagation and In Vitro Propagation of Ceropegia Species6.1 Introduction
6.2 Propagation of Ceropegia
6.2.1 Growing Conditions
6.2.2 Propagation
6.2.3 Grower's Tips
6.3 Seed Dormancy
6.4 In Vitro Propagation
6.4.1 Ceropegia attenuata
6.4.2 Ceropegia bulbosa
6.4.3 Ceropegia candelabrum
6.4.4 Ceropegia elegans
6.4.5 Ceropegia ensifolia
6.4.6 Ceropegia evansii
6.4.7 Ceropegia fantastica
6.4.8 Ceropegia fimbriifera
6.4.9 Ceropegia hirsuta
6.4.10 Ceropegia intermedia
6.4.11 Ceropegia jainii
6.4.12 Ceropegia juncea
6.4.13 Ceropegia lawii
6.4.14 Ceropegia maccannii 6.4.15 Ceropegia mahabalei6.4.16 Ceropegia media
6.4.17 Ceropegia noorjahaniae
6.4.18 Ceropegia oculata
6.4.19 Ceropegia odorata
6.4.20 Ceropegia panchganiensis
6.4.21 Ceropegia pullaiahii
6.4.22 Ceropegia pusilla
6.4.23 Ceropegia rollae
6.4.24 Ceropegia sahyadrica
6.4.25 Ceropegia santapaui
6.4.26 Ceropegia spiralis
6.4.27 Ceropegia thwaitesii
6.4.28 Ceropegia woodii
6.5 Synthetic Seed Technology
6.6 In Vitro Tuberization
References
7: Molecular Taxonomy of Indian Brachystelma and Ceropegia
7.1 Introduction
7.2 Tribe Ceropegiae in India

Citation preview

Monograph on Brachystelma and Ceropegia in India

Monograph on Brachystelma and Ceropegia in India

Thammineni Pullaiah Subbiah Karuppusamy Kondragunta Sri Rama Murthy

CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 © 2019 by Taylor & Francis Group, LLC CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works Printed on acid-free paper International Standard Book Number-13: 978-0-3671-8381-3 (Hardback) This book contains information obtained from authentic and highly regarded sources. Reasonable efforts have been made to publish reliable data and information, but the author and publisher cannot assume responsibility for the validity of all materials or the consequences of their use. The authors and publishers have attempted to trace the copyright holders of all material reproduced in this publication and apologize to copyright holders if permission to publish in this form has not been obtained. If any copyright material has not been acknowledged please write and let us know so we may rectify in any future reprint. Except as permitted under U.S. Copyright Law, no part of this book may be reprinted, reproduced, transmitted, or utilized in any form by any electronic, mechanical, or other means, now known or hereafter invented, including photocopying, microfilming, and recording, or in any information storage or retrieval system, without written permission from the publishers. For permission to photocopy or use material electronically from this work, please access www. copyright.com (http://www.copyright.com/) or contact the Copyright Clearance Center, Inc. (CCC), 222 Rosewood Drive, Danvers, MA 01923, 978-750-8400. CCC is a not-for-profit organization that provides licenses and registration for a variety of users. For organizations that have been granted a photocopy license by the CCC, a separate system of payment has been arranged. Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. Library of Congress Cataloging‑in‑Publication Data Names: Pullaiah, T., author. Title: Monograph on Brachystelma and Ceropegia in India / authors: Thammineni Pullaiah, Subbiah Karuppusamy, Kondragunta Sri Rama Murthy. Description: Boca Raton, FL : CRC Press, Taylor & Francis Group, 2019. | Includes bibliographical references and index. Identifiers: LCCN 2019006778 | ISBN 9780367183813 (hardback : alk. paper) | ISBN 9780429061233 (ebook) Subjects: LCSH: Brachystelma--India. | Ceropegia--India. Classification: LCC QK495.A815 M66 2019 | DDC 583/.930954--dc23 LC record available at https://lccn.loc.gov/2019006778 Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com

Contents Preface.........................................................................................................................ix Authors.........................................................................................................................xi 1. Introduction......................................................................................................... 1 1.1 Introduction................................................................................................ 1 References............................................................................................................. 2 2. Brachystelma Taxonomy..................................................................................... 5 2.1 Introduction................................................................................................ 5 2.2 Systematic Enumeration............................................................................ 8 References........................................................................................................... 78 3. Traditional Uses, Pharmacognostic, Phytochemical, Pharmacological, and In Vitro Propagation Studies in Brachystelma Species............................81 3.1 Introduction.............................................................................................. 81 3.2 Edible Uses.............................................................................................. 81 3.3 Medicinal Uses........................................................................................ 83 3.4 Pharmacognostical Studies......................................................................84 3.5 Phytochemical Studies............................................................................. 85 3.5.1 Antioxidant Activity.................................................................. 86 3.5.2 Nutritional and Mineral Studies................................................ 87 3.5.3 Bromatological Analysis............................................................ 88 3.6 In Vitro Propagation Studies.................................................................... 88 3.6.1 In Vitro Propagation Studies of Brachystelma glabrum........... 89 3.6.2  In Vitro Propagation Studies of B. ngomense, B. pulchellum, and B. pygmaeum���������������������������������������������90 3.7 Conclusion................................................................................................ 94 References........................................................................................................... 94 4. Ceropegia Taxonomy.........................................................................................97 4.1 Introduction..............................................................................................97 4.1.1 Botanical History of Indian Ceropegia.....................................97 4.2 Systematic Enumeration..........................................................................99 References......................................................................................................... 214 5. Traditional Uses, Phytochemistry, and Pharmacology of Ceropegia Species......221 5.1 Introduction............................................................................................ 221 5.2 Ornamental Uses.................................................................................... 221 5.3 Traditional Uses..................................................................................... 222 5.3.1 Edible Uses.............................................................................. 222 5.3.2 Traditional Medicinal Uses...................................................... 222 v

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Contents 5.4

Pharmacognostic Studies of Ceropegia Species................................... 229 5.4.1 Powder Characteristics............................................................. 230 5.4.1.1 Physicochemical Analysis....................................... 230 5.5 Phytochemistry of Ceropegia Species................................................... 231 5.5.1 Chemical Constituents in Ceropegia....................................... 231 5.5.2 Major Bioactive Constituents of Ceropegia Species............... 232 5.5.3 Chemical Synthesis of Cerpegin.............................................. 243 5.6 Pharmacology of Ceropegia Species.....................................................244 5.6.1 Antimicrobial Activity.............................................................244 5.6.2 Antiurolithic Activity............................................................... 245 5.6.3 Analgesic Activity....................................................................246 5.6.4 Hepatoprotective and Antioxidant Activity.............................246 5.6.5 Antioxidant Activity................................................................246 5.6.6 Anticancer Activity.................................................................. 247 5.6.7 Antidiabetic Activity................................................................ 247 5.6.8 Anti-inflammatory Activity.....................................................248 5.6.9 Gastroprotective Effect............................................................248 5.6.10 Inhibitory Activity against Peptidases.....................................248 References......................................................................................................... 249 6. Propagation and In Vitro Propagation of Ceropegia Species...................... 255 6.1 Introduction............................................................................................ 255 6.2 Propagation of Ceropegia...................................................................... 255 6.2.1 Growing Conditions................................................................. 255 6.2.2 Propagation.............................................................................. 255 6.2.3 Grower’s Tips........................................................................... 256 6.3 Seed Dormancy...................................................................................... 256 6.4 In Vitro Propagation............................................................................... 257 6.4.1 Ceropegia attenuata................................................................264 6.4.2 Ceropegia bulbosa................................................................... 265 6.4.3 Ceropegia candelabrum.......................................................... 269 6.4.4 Ceropegia elegans.................................................................... 269 6.4.5 Ceropegia ensifolia.................................................................. 271 6.4.6 Ceropegia evansii.................................................................... 271 6.4.7 Ceropegia fantastica................................................................ 271 6.4.8 Ceropegia fimbriifera............................................................... 273 6.4.9 Ceropegia hirsuta.................................................................... 273 6.4.10 Ceropegia intermedia.............................................................. 273 6.4.11 Ceropegia jainii....................................................................... 274 6.4.12 Ceropegia juncea..................................................................... 275 6.4.13 Ceropegia lawii........................................................................ 276 6.4.14 Ceropegia maccannii............................................................... 277 6.4.15 Ceropegia mahabalei............................................................... 277 6.4.16 Ceropegia media...................................................................... 277 6.4.17 Ceropegia noorjahaniae.......................................................... 278 6.4.18 Ceropegia oculata.................................................................... 278 6.4.19 Ceropegia odorata................................................................... 279 6.4.20 Ceropegia panchganiensis....................................................... 279

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6.4.21 Ceropegia pullaiahii................................................................ 279 6.4.22 Ceropegia pusilla..................................................................... 281 6.4.23 Ceropegia rollae...................................................................... 281 6.4.24 Ceropegia sahyadrica.............................................................. 283 6.4.25 Ceropegia santapaui................................................................ 283 6.4.26 Ceropegia spiralis.................................................................... 283 6.4.27 Ceropegia thwaitesii................................................................ 286 6.4.28 Ceropegia woodii..................................................................... 286 6.5 Synthetic Seed Technology.................................................................... 286 6.6 In Vitro Tuberization.............................................................................. 288 References......................................................................................................... 290 7. Molecular Taxonomy of Indian Brachystelma and Ceropegia.................... 295 7.1 Introduction............................................................................................ 295 7.2 Tribe Ceropegiae in India...................................................................... 296 7.3 Molecular Markers Used for Phylogenetic Analysis of Tribe Ceropegiae�������������������������������������������������������������������������������������������� 297 7.3.1 Molecular Relationship of Indian Brachystelma and Ceropegia................................................................................. 297 7.3.2 ITS Phylogeny.......................................................................... 298 7.3.3 cpDNA Phylogeny.................................................................... 298 7.3.4 The Combined of ITS and cpDNA Phylogeny........................ 301 7.3.5 Recent Phylogenetic Radiation in Brachystelma and Ceropegia.................................................................................302 7.3.6 Diagnosis of New Species Through Molecular Systematic.....302 7.3.6.1 Brachystelma mahajanii and B. vartakii................302 7.3.6.2 Ceropegia mizoramensis and Ceropegia murlensis.....304 7.3.6.3 Brachystelma mahendragiriensis............................306 7.4 Problems in Phylogenetic Studies..........................................................306 7.5 Conclusion..............................................................................................306 References.........................................................................................................306 8. Pollination Biology of Ceropegia and Brachystelma Species.......................309 8.1 Introduction............................................................................................309 8.2 Self-Pollination......................................................................................309 8.3 Pollinators.............................................................................................. 311 8.4 Pollinators Diversity.............................................................................. 311 8.5 Pollinator Specificity Through Floral Scent.......................................... 312 8.6 Pollination Mechanism in Ceropegias................................................... 314 8.7 Pollination Systems in Ceropegias and Its Phylogenetic Relationships.....315 8.8 Pollination System in Ceropegia juncea............................................... 319 8.9 Conclusion.............................................................................................. 321 References......................................................................................................... 322 9. Threats and Conservation of Brachystelma and Ceropegia in India......... 325 9.1 Introduction............................................................................................ 325 9.2 Trade and CITES................................................................................... 325 9.3 Possible Threats..................................................................................... 326

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Contents 9.4 Collection for Ornamentals................................................................... 327 9.5 Pollination Factors................................................................................. 328 9.6 Seed Setting and Germination Efficiency............................................. 328 9.7 Natural Calamities................................................................................. 328 9.8 Biotechnological Approach on Conservation........................................ 329 9.9 Problems in Conservation...................................................................... 329 References......................................................................................................... 329

10. Contributors for the Taxonomy of Indian Brachystelma and Ceropegia......333 Von Rheede (1636–1691).................................................................................. 333 Carolus Linnaeus (1707–1778)......................................................................... 333 William Roxburgh (1751–1815)....................................................................... 334 Nathaniel Wallich (1786–1854)........................................................................ 334 Robert Wight (1796–1872)................................................................................ 335 Robert Graham (1805–1839)............................................................................ 336 Joseph Decaisne (1807–1882)........................................................................... 336 Colonel Richard Henry Beddome (1830–1911)................................................ 336 Sir Joseph Dalton Hooker (1817–1911)............................................................ 337 James Sykes Gamble (1847–1925)................................................................... 337 Herbert Franz Josef Huber (1931–2005)........................................................... 338 Anis Ahmad Ansari (1953–).............................................................................. 339 Prof. Peter Bruyns (1957–)................................................................................ 339 Prof. Srirang Ramchandra Yadav (1954–).........................................................340 Dr. Ulrich Meve (1958–)...................................................................................340 Dr. Kothareddy Prasad (1985–)......................................................................... 341 Index of Scientific Names�������������������������������������������������������������������������������������� 343

Preface In recent years, many new species of Brachystelma and Ceropegia have been described from India. These two genera have not been revised in India and hence an attempt has been made to bring out a consolidated account on Brachystelma and Ceropegia in India along with other aspects of these two genera. We thank the editors of Taiwania, Bangladesh Journal of Plant Taxonomy, Nordic Journal of Botany, Journal of Threatened Taxa, and Pleione for giving permission to reproduce the figures. We thank authorities of the Central National Herbarium; Botanical Survey of India, Western Circle; Botanical Survey of India, Southern Circle; Botanical Survey of India, Deccan Regional Centre; Agarkar Research Institute, Pune; Shivaji University, Kolhapur; Trans Disciplinary University (FRLHT), Bangalore; Rapinat Herbarium, Tiruchirappalli, and Blatter Herbarium, Mumbai, for giving permission to consult the herbaria and their libraries. Dr. K. Raja Kullayiswamy has drawn several diagrams and shared many of his photographs for this book. He also collected and gave description of Brachystelma volubile. The authors are grateful for all the help and service rendered by him. We thank Dr. Kanchi N. Gandhi of Harvard University for valuable suggestions. Prof. Vijayasankar Raman of the University of Mississippi has critically reviewed our book proposal and has gone through the manuscript on Brachystelma and gave many suggestions and comments for which the authors are grateful to him. We also thank Prof. van Staden from South Africa, Dr. Signe Nybø from Norway, Dr. Andrew E. Whittington from the UK, and an anonymous reviewer for critical review of our book proposal and for their valuable suggestions.

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Authors

Thammineni Pullaiah, PhD, is a former professor at the Department of Botany at Sri Krishnadevaraya University in Andhra Pradesh, India, where he has taught for more than 35 years. He has held several positions at the university, including dean, faculty of Biosciences, head of the Department of Botany, head of the Department of Biotechnology, and member of the Academic Senate. He was president of the Indian Botanical Society (2014), president of the Indian Association for Angiosperm Taxonomy (2013), and Fellow of Andhra Pradesh Akademi of Sciences. He was awarded the Panchanan Maheshwari Gold Medal, the Dr. G. Panigrahi Memorial Lecture Award of the Indian Botanical Society, the Prof. Y.D. Tyagi Gold Medal of the Indian Association for Angiosperm Taxonomy, and a Best Teacher Award from the Government of Andhra Pradesh. Under his guidance, 54 students obtained their doctoral degrees. He has authored 51 books, edited 18 books, and published over 330 research papers, including reviews and book chapters. His books include Ethnobotany of India (5 volumes published by Apple Academic Press), Flora of Andhra Pradesh (5 volumes), Flora of Eastern Ghats (4 volumes), Flora of Telangana (3 volumes), Encyclopaedia of World Medicinal Plants (5 volumes, 2nd edition), and Encyclopaedia of Herbal Antioxidants (3 volumes). He was also a member of the Species Survival Commission of the International Union for Conservation of Nature (IUCN). Professor Pullaiah received his PhD from Andhra University, India, attended Moscow State University, Russia, and worked as Postdoctoral Fellow during 1976–1978. Dr. Subbiah Karuppusamy, PhD, obtained his MSc degree from Madurai Kamaraj University, Madurai, and PhD degree from Gandhigram Rural Institute, Deemed University, Tamil Nadu. Later on, he worked as a Postdoctoral Fellow in the Department of Biotechnology and a research associate in the Department of Botany, Sri Krishnadevaraya University, Anantapur. He has published more than 120 research papers in reputed journals and coauthored five books—Flora of Eastern Ghats (Volumes 3 and 4), Flora of Andhra Pradesh (Volume 5), Revised Flora of Andhra Pradesh (Volume 3), and Caralluma (Sensu lato). He is the recipient of the Prof. Y.S. Murthy Gold xi

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Medal from the Indian Botanical Society and the Rolla S. Rao award from the Indian Association for Angiosperm Taxonomy. He is a Fellow of the Indian Association for Angiosperm Taxonomy and the Indian Botanical Society. He is the principal investigator of three major research projects funded by the UGC, SERB, and NIF. Under his guidance, four students obtained their doctoral degrees. Dr. Kondragunta Sri Rama Murthy received his master’s degree (1994) in botany from Acharya Nagarjuna University and thereafter obtained his PhD, (1998) from Sri Krishnadevaraya University, Anantapur. He was awarded the DST-SERC Young Scientist Project (2004–2006) in the field of domestication of wild relatives of cultivated legumes for food and agriculture. He worked as professor and head of the Department of Botany, Andhra Loyola College, Vijayawada, for four years and professor and head of the Department of Biotechnology for eight years in Montessori Mahila Kalasala, Vijayawada. He is the principal investigator for two UGC and two CSIR projects and completed them successfully. Under his guidance, six students were awarded doctoral degrees, and seven students received their master of philosophy degrees. He is the coauthor of Flora of Andhra Pradesh (Volume 2) and Flora of Eastern Ghats (Volumes 2 and 3). He has published 110 scientific research papers in nationally and internationally reputed journals and also filed five Indian patents. He is a life member of the International Society of Plant Morphologists and the Association of Biotechnology and Pharmacy.

1 Introduction

1.1 Introduction Apocynaceae, including its subfamilies, has about 4800 species and 480 genera (Struwe et al., 1994; Mabberley, 2008). Endress and Bruyns (2000) proposed a revised classification of Apocynaceae, which includes five subfamilies: Rauvolfiodeae, Apocynoideae, Periplocoideae, Secamonoideae, and Asclepiadoideae. The subfamily Asclepiadoideae was formerly known as Asclepiadaceae (Endress and Bruyns, 2000). Asclepiadoideae has 177 genera and 3,000 species (Meve, 2002) and has four tribes – Fockeeae, Asclepiadeae, Marsdenieae, and Ceropegieae (Endress and Bruyns, 2000). Asclepiadoideae has the most elaborate and complex flowers of all dicots (Endress, 1994). The genera Brachystelma R. Br. ex Sims. and Ceropegia L. were earlier included in the family Asclepiadaceae. In APGIV Classification, the former family Asclepiadaceae has been reduced to subfamily Asclepiadoideae under the family Apocynaceae, and these two genera have been included in tribe Ceropegieae, subfamily Asclepiadoideae. The roughly 710 species of the tribe Ceropegieae (Apocynaceae-Asclepiadoideae) are made up of the four major groups: Brachystelma (120 spp.), Ceropegia (200 spp.), the stem-succulents popularly known as the stapeliads (350 spp. in 33 genera), and an early-divergent group of about 45 species (Bruyns et al., 2014, 2015). The genus Brachystelma was described by Robert Brown in 1822 based on Meerburgh’s Stapelia tuberosa from the Cape of Good Hope, South Africa. It is more similar vegetatively to the genus Ceropegia (erect/twining leafy herbs with terete stems) than to Frerea (pendulous, succulent, leafy herbs with terete stems) or Caralluma R. Br. (leafless herbs with fleshy four-angled stems) of the same tribe Ceropegieae. Brachystelma has distinctive features from others with slender stems (stout and fleshy in Frerea), corolla not tubular (tubular in Ceropegia), and relatively narrow lobes (compared to Ceropegia). The genus ranks second (in number of species) in the tribe Ceropegieae and is represented by about 160 species distributed chiefly in sub-Saharan Africa, India, Sri Lanka, Southeast Asia, and Northern Australia (Meve, 2002; Venu and Prasad, 2015) but missing in Arabia (Meve, 2002; Masinde, 2007; Mabberley, 2008). The genus Ceropegia Linnaeus (1753: 211), the largest genus in the tribe Ceropegieae (Asclepiadoideae, Apocynaceae), is represented by 244 taxa worldwide (cf., The Plant List, 2013), distributed only in the Old World ranging from the Spanish Canary Islands in the West, through central, southern, and northern Africa, Madagascar, Arabia, India, Southeastern Asia and Southwestern Pacific region (including Papua New Guinea, Indonesia, and Philippines), and Northeastern Australia (Ansari, 1984; 1

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Monograph on Brachystelma and Ceropegia in India

Meve and Liede-Schumann, 2007; Murthy et al., 2012; Bruyns et al., 2015). Ceropegia species are found in subtropical Africa toward the eastern side of the continent (Dyer, 1983; Bruyns, 2003; Bruyns et al., 2015). Significant lantern flower diversity is also found in Madgascar and in the Indian subcontinent (Bruyns, 2003; Murthy et al., 2012; Bruyns et al., 2015). Ceropegia spp. have certain universally used names, viz., lantern flower, Christensen parasol flower, parachute flower, bushman’s pipe, string of hearts, snake creeper, wine-glass vine, rosary vine, necklace vine flower, Chinese lantern, lantern plant, trap flowers, and pitfall trap flowers (Yadav, 1996; Quattrocchi, 2000). The maximum diversity of Ceropegia is found in subtropical Africa toward the eastern side of the continent where more than 50 species have been reported (Dyer, 1983). Apart from this region, significant species diversity occurs in Madagascar and in the Indian subcontinent (Bruyns, 1997). In China, there are 17 species with 2 species overlapping with India (Li et al., 1995). In India, there seems to be two major distributions of this genus, the Himalayan region and Peninsular region. The Himalayan species do not possess tubers and are non-succulent and herbaceous (Bruyns, 1997). Ceropegia was revised by Huber (1957), who recognized 153 species in 21 sections, including 1 for the 6 species of Riocreuxia, which he included in Ceropegia. Apart from Huber’s placing of Riocreuxia in Ceropegia (from where it was removed again by Dyer, 1980), the generic circumscription of Ceropegia has not been controversial, as the generic synonymy of Ceropegia shows (Huber, 1957; Dyer, 1980). Huber’s sections (Huber, 1957) have not been used by any subsequent authors, with Dyer (1980) dismissing Huber’s emphasis on pubescence as having led to ‘some unnatural groupings’. Exploration since 1957 has unearthed much new information, from which regional revisions were published by Ansari (1984) for India; Bruyns (1985,  1986, 1989) for the former Cape Province in South Africa, the Canary Islands, the Arabian Peninsula, and Namibia, respectively; Dyer (1980, 1983) for Southern Africa; Archer (1992) and Masinde (2012) for East Africa; and Forster (1988) for the single Australian species. Many new species of Brachystelma and Ceropegia have recently been described from India. These two genera have not been revised in India and hence an attempt has been made to bring out a consolidated account on Brachystelma and Ceropegia in India along with other aspects of these two genera.

REFERENCES Angiosperm Phylogeny Group IV (APGIV). 2016. An update of the Angiosperm Phylo­ geny Group classification for the orders and families of flowering plants APG IV. Bot. J. Linn. Soc. 181: 1–20. Ansari, M.Y. 1984. Asclepiadaceae: Genus–Ceropegia. Fascicles of Flora of India, Fascicle 16. Botanical Survey of India, Howrah, India, pp. 1–35. Archer, P.G. 1992. Kenya Ceropegia Scrapbook. Notes and Records of Some Kenya Ceropegia. Artemis, Hobart, Australia. Bruyns, P.V. 1985. Notes on ceropegias of the Cape Province. Bradleya 3: 1–47. Bruyns, P.V. 1986. The genus Ceropegia on the Canary Islands (AsclepiadaceaeCeropegieae). Beitr. Biol. Pflanz. 60: 427–458. Bruyns, P.V. 1989. Studies in the flora of Arabia XXIV: The genus Ceropegia in Arabia. Notes Royal Bot. Gard., Edinburgh 45: 287–326.

Introduction

3

Bruyns, P.V. 1997. A note on Ceropegia L. (Asclepiadaceae) of Silent Valley, Kerala, India. Rheedea 7(2): 107–114. Bruyns, P.V. 2003. Three new succulent species of Apocynaceae (Asclepiadoideae) from Southern Africa. Kew Bull. 58: 427–435. Bruyns, P.V., C. Klak, and P. Hanáček. 2014. Evolution of the stapeliads (Apocynaceae– Asclepiadoideae) – Repeated major radiation across Africa in an Old World group. Mol. Phylogen. Evol. 77: 251–263. Bruyns, P.V., C. Klak, and P. Hanáček. 2015. Recent radiation of Brachystelma and Ceropegia (Apocynaceae) across the Old World against a background climate change. Mol. Phylogen. Evol. 90: 49–66. Dyer, R.A. 1980. Brachystelma, Ceropegia, Riocreuxia. In: Leistner, O.A. (Ed.), Flora of Southern Africa, Botanical Research Institute, Pretoria, South Africa, Vol. 27, No. 4, pp. 1–91. Dyer, R.A. 1983. Ceropegia, Brachystelma and Riocreuxia in Southern Africa. Balkema, Rotterdam, the Netherlands. Endress, M.E. and P.V. Bruyns. 2000. A revised classification of the Apocynaceae s.l.Bot. Rev. 66: 1–56. Endress, P.K. 1994. Diversity and Evolutionary Biology of Tropical Flowers. University Press, Cambridge, UK. Forster, P.I. 1988. Studies on the Australiasian Asclepiadaceae. I. Brachystelma Sims in Australia. Nuytsia 6: 285–294. Huber, H. 1957. Revision der Gattung Ceropegia. Mem. Soc. Broter. 12: 1–203. Li, P.T., M.G. Gilbert, and W.D. Stevens. 1995. Asclepiadaceae. In: Wu, Z.Y. and P.H. Raven (Eds.), Flora of China, Science Press, Beijing, China, Vol. 16, pp. 189–270. Mabberley, D.J. 2008. Mabberley’s Plant-book: A Portable Dictionary of Plants, Their Classification and Uses. Cambridge University Press, Cambridge, UK. Masinde, P.S. 2007. A revision of Brachystelma Sims (Apocynaceae: AsclepiadoideaeCeropegieae) in East Africa. Kew Bull. 62: 37–84. Masinde, P.S. 2012. Ceropegia. In: Beentje, H. (Ed.), Flora of Tropical East Africa, Apocynaceae (Part 2), Royal Botanic Gardens, Kew, pp. 220–291. Meve, U. 2002. Brachystelma. In: Albers, F. and U. Meve (Eds.), Illustrated Handbook of Succulent Plants: Asclepiadaceae, Springer, New York, pp. 20–46. Meve, U. and S. Liede-Schumann. 2007. Ceropegia (Apocynaceae, Ceropegieae, Stapeliinae): Paraphyletic, but still taxonomically sound. Ann. Missouri Bot. Gard. 94: 392–406. Murthy, K.S.R., R. Kondamudi, M.C. Reddy, S. Karuppusamy, and T. Pullaiah. 2012. Check-list and conservation strategies of the genus Ceropegia in India. Int. J. Biodiv. Conserv. 4: 304–315. Quattrocchi, U. 2000. CRC World Dictionary of Plant Names – Common Names, Scientific Names, Eponyms, Synonyms and Etymology. CRC Press, Boca Raton, FL. Struwe, L., V.A. Albert, and B. Bremer. 1994. Cladistics and family level classification of the Gentianales. Cladistics 10: 175–206. Venu, P. and K. Prasad. 2015. The existential crisis in Indian Brachystelmas (Apocynaceae). Curr. Sci. 109(4): 680–682. Yadav, S.R. 1996. Fly trap flowers of the Western Ghats. Hornbill 1: 2–7.

2 Brachystelma Taxonomy

2.1 Introduction The genus Brachystelma belongs to the tribe Ceropegieae of the subfamily Asclepiodoideae of the family Apocynaceae. The name Brachystelma comes from the Greek words “brachy” (short) and “stelma” (crown), a description of the short corona in the flower of some species. The genus Brachystelma was described by Robert Brown in 1822 based on Meerburgh’s Stapelia tuberosa from the Cape of Good Hope, South Africa. It is more similar vegetatively to the genus Ceropegia (erect/twining leafy herbs with terete stems) than to Frerea (pendulous, succulent, leafy herbs with terete stems) or Caralluma R. Br. (leafless herbs with fleshy fourangled stems) of the same tribe Ceropegieae. Brachystelma has distinctive features from others with slender stems (stout and fleshy in Frerea), corolla not tubular (tubular in Ceropegia) and the lobes relatively narrow (compared to Ceropegia). The members of Brachystelma are essentially herbaceous, bearing medium-sized tubers and fleshy roots, stems terete, unbranched to 1.5  m high (a couple of them are twiners) with opposite leaves; variedly ornamented flowers, either solitary or few (more than 20 flowers in B. swarupa), usually in umbels, free calyx lobes, corolla tube absent, rarely much smaller (B. brevitubulatum and B. parviflorum); lobes usually broad (filiform in B. attenuatum), glabrous (B. elenaduense and B. mahajanii), hairy (B. laevigatum, B. vartakii, and B.naorojii) and with striations (B. nallamalayana), gorgeously colored (brown/black with pink hairs in B. malwanense; white to whitishpink in B. vartakii; deep purple in B. elenaduense; greenish yellow in B. mahajanii), either spreading and star-like (B. penchalakonense and B.pullaiahii), rarely reflexed and the whole flower resembling the Ashoka emblem (B. ciliatum) or fused at the tips to give the appearance of a bird’s cage (B. brevitubulatum, B. kolarense, and B. malwanense), and corona uniquely structured and brilliantly colored (yellow in B. pullaiahii, greenish or reddish-yellow in B. penchalakonense, black in B. nallamalayana, pinkish in B. vartakii, greenish yellow spotted purple in B. mahajanii), biseriate, outer cup-like, either annular or angled, shallowly undulate or five-lobed (which is often further lobed), inner forming a cup or not, with five simple oblong segments, incumbent over stigma; pollen masses solitary in each locule, with extra pellucid margins on the inner side of pollinia. Follicles are solitary or in pairs, linear, bearing comatose seeds. On the whole, with brilliant display of corolla and coronal structures, they appear small but beautiful (Venu and Prasad, 2015). The first account of Indian Brachystelma was presented by Hooker, enumerating seven of them discovered by him; four species from peninsular India, three of them based on Beddome’s collections (B. glabrum, B. brevitubulatum, and B. volubile) 5

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and one of Law’s collection (B. maculatum); two from northwestern India based on Royle’s collections (B. parviflorum and B. attenuatum) and one from sub-Himalaya (Uttar Pradesh) based on Hamilton’s collection (B. laevigatum). All these species exhibit either erect (five species) or twining habit (two species). They show highly restricted distribution and are yet to be collected outside the type localities. Hooker had stated that his specific descriptions were “imperfect” because these species have very complex coronal processes and he faced difficulties in the description of the colors as well as coronal structures of the flowers in dried specimens. He assumed that the future explorers, with newer collections and dissections on fresh materials, would possibly enrich diagnostics and species delimitations. Hooker (1883) in his Flora of British India (volume 4, page 65) rightly pointed out that there might be many species of this singular genus in India, which owing to their slender habit and inconspicuous flowers may be overlooked. As predicted by Hooker, later published floras added a few more new species, B. bourneae and B. rangacharii by Gamble (1921) from Madras Presidency and B. pauciflorum Duthie (1911) from the Bahariach district in Uttar Pradesh. Thus, of the 10 species listed until the 1930s, excluding 2, which have been claimed to be recollected—B. volubile from Kadapa hills, Andhra Pradesh (Kullayiswamy et  al., 2012) and B. brevitubulatum from the Tiruvannamalai district, Tamil Nadu (Vijayasankar et al., 2003)—others remained hidden and evaded recollections by the explorers. Unfortunately, the latter Floras (Sharma et al., 1984; Srinivasan et al., 1987; Hanumanthappa, 1997) merely cited these species based on the old, often single collections cited by earlier workers. Even the generic review was done based on old collections in the revision of Asclepiadaceae (Jagtap and Singh, 1999), with no additional data. It is pertinent to mention that many species are devoid of description on intricate details regarding coronal structures and fruits/seeds crucial for better delimitation of species and perhaps the genus itself. B. attenuatum and B. parviflorum are based on mere illustrations (both in Wight herbarium) and without any specimens; an illustration along with a single collection as old as 200 years is the basis of description for B. laevigatum; B. pauciflorum has neither a specimen nor an illustration for the basis of the name. We have nothing but representation of types in the case of B. bourneae, B. glabrum, B. maculatum, and B. rangacharii. This has in fact forced reconsideration on their continued existence in India (Venu and Prasad, 2015). There were no additions in Indian Brachystelma for almost 40 years (1930–1970), but altogether 13 taxa were added between 1970 and 2015 and another 7 species in the year 2016 alone (Table 2.1). It is unfortunate that except for B. ciliatum, other names are associated with single collections and single reports. Till date, 33 species have been reported from India (majority from Peninsular India (Table 2.1); 31 of them are endemic and 30 (about 94%) had no recollections after the types (Venu and Prasad, 2015; Prasad et al., 2016, 2017). In the Botanical Survey of India, Pune Herbarium, apart from type specimens, there is only one specimen of Brachystelma. In Madras Herbarium (apart from type specimens), there are only 7 specimens of Brachystelma to which 3 specimens have been added recently. Brachystelma laevigatum has no type specimen. Even a description as mentioned by Duthie is based on only one specimen. For many species there is only flower description and no description of fruit, e.g., B. penchalakonense. When asked about the situation, authors of this species informed us that they could collect only two specimens. The same is the case for many other species. The majority of the species occur in small

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TABLE 2.1 List of New Species/New Records of Brachystelma Species from India Between 1970 and 2017 1971–1980

1. B. elenaduense Char, Proc. Indian Sci. Congr. Assoc., 1971, 58, 435 ex Sathyan, Curr. Sci., 1978, 47(24), 965. 1981–1990

2. B. ciliatum Arekal & T.M. Ramakrishna, Curr. Sci., 1981, 50, 145. 3. B. kolarense Arekal & T.M. Ramakrishna, Proc. Indian Acad. Sci., Plant Sci., 1981, 90, 203. 1990–2001

4. B. malwanense S.R. Yadav & N.P. Singh, Kew Bull., 1993, 48, 59. 5. B. naorojii P. Tetali et al., Rheedea, 1998, 8, 75. 6. B. swarupa K.K. Kumar & Goyder, Kew Bull., 2001, 56, 210. 2011–2018

7. B. pullaiahii B.R.P. Rao et al., Taiwania, 2011, 56(3), 223. 8. B. nallamalayana K. Prasad & B.R.P. Rao, J. Threat. Taxa, 2013, 5(14), 4904. 9. B. penchalakonense Rasingam et al., Kew Bull., 2013, 68, 663. 10. B. mahajanii Kambale & S.R. Yadav, Kew Bull., 2014, 69(1), 9493 (2). 11. B. vartakii Kambale & S.R. Yadav, Kew Bull., 2014, 69(1), 9493 (4). 12. B. annamacharyae Prasad et al., Nord. J. Bot., 2015, 34, 360. 13. B. seshachalamense Prasad & Prasanna, Bangladesh J. Plant Taxon., 2016, 23, 53. 14. B. matthewianum Bruyns & Britto, Haseltonia, 2016, 22, 48, 51. 15. B. rapinatianum Bruyns & Britto, Haseltonia, 2016, 22, 48, 50. 16. B. saldanhae Bruyns & Britto, Haseltonia, 2016, 22, 48, 52. 17. B. gondwanense Govekar et al., Rheedea, 2016, 26, 145. 18. B. nigidianum Raja Kullayisw et al., Kew Bull., 2016, 71, 1. 19. B. shrirangii Kambale et al., Rheedea, 2016, 26, 145. 20. B. mahendragiriense Prasad et al., Rheedea, 2017. 27, 135. 21. B. ananathapuramense Prasad et al., Kew Bull., 2018. 73, 16. 22. B. vemanae A. Madhusudhana Reddy, M.V. Suresh Babu & K. Prasad, Nordic J. Bot. 2018. 36, 1. areas with localized populations. There is also a chance that many species might be evading collections because they are small, tender, and delicate with short growing/ flowering periods, and grassier than any distinctive appearance in grasslands/grassdominated habitats. It is unfortunate that grasslands/grass-dominated habitats, where its members often thrive, are regarded as waste and unproductive with no worthwhile protection. They are encroached upon for agriculture and often fragmented by habitations, urbanization, forest fires, and invasive species. The habitats are also prone to elimination since many grass species, primarily those of Cymbopogon, are harvested as fodder or for thatching by locals. There is the likelihood of these species being removed along with them. The edible tubers of the members of this genus and allied genus Ceropegia, known variedly as Nematai, Nematigaddalu, Potha

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Jougu Nimatayalu, Petta Jougu Nimatayalu, Singati galya, are often dug out by the locals. Wild animals, rodents, wild boar, and langurs also relish these tubers and thus threaten their natural regeneration. Many focused publications on endemic and threatened species of India surprisingly omitted this genus as a whole (Ahmedullah and Nayar, 1987; Nayar and Sastry, 1987–1990). Rao et al. (2003) and Nayar (1996) placed B. bourneae under “indeterminate” and “possibly extinct” categories, respectively. In spite of their rarity and distinctiveness from African Brachystelma (Meve, 2007; Surveswaran et al., 2009), they were neither considered for any focused collection nor regarded for IUCN conservation status. The genus as a whole falls in the most deserving zone for focused conservation. The whole group is to be attempted in a project mode primarily for recollections of all the documented species, taxonomic revision, assessing their distribution in the field and assigning IUCN conservation status, promoting studies on coronal structures, pollination ecology, ex situ conservation, and rehabilitating them in protected natural habitats by acclimatizing and multiplying them in gardens, thereby giving a whole insight into their taxonomy and conservation. They are true blinking stars of angiosperms and require due care. In India, Brachystelma is represented by 33 species and, 31 of them are endemic to the country (Venu and Prasad, 2015; Prasad et al., 2016, 2017, 2018). Duthie (1911) reported two species of Brachystelma from the Upper Gangetic Plain. Gamble (1923) described six species of Brachystelma from Presidency of Madras. S.R. Srinivasan (1987) reported four species of Brachystelma from the state of Tamil Nadu. Ramakrishna et al. (1995) reported seven species of Brachystelma from the state of Karnataka. Jagtap and Singh (1999) in Fascicles of Flora India reported 14 species of Brachystelma from India. Karthikeyan et al. (2009) listed 17 species of Brachystelma in India. Pullaiah et al. (2011) reported three species of Brachystelma from Eastern Ghats. The genus is now represented by 9 species in Eastern Ghats (Hooker, 1883; Gamble, 1921; Rao et al., 2011; Prasad and Rao, 2013; Rasingam et al., 2013; Prasad et al., 2017). Four species are with twining habit, namely Brachystelma brevitubulatum Hook. f., B. nigidianum Kullayis., B. seshachalmensis, and B. volubile Hook. f., which are endemic to Tamil Nadu and Andhra Pradesh, respectively. These species were recollected by Kullayiswamy et al. (2016) (B. volubile, Tirumala Hills, Andhra Pradesh) and Vijayasankar et al. (2003) (B. brevitubulatum, Tiruvannamalai District, Tamil Nadu). Singh et al. (2015) reported that 16 species of Brachystelma are endemic to India. Sasidharan (2004) recorded only one species of Brachystelma (B. swarupa) from Kerala.

2.2  Systematic Enumeration Brachystelma R.Br., Bot. Mag. tab. 2343. 1822. nom. cons. Herbs erect or twining, perennial. Rootstock usually a subglobose tuber (sometimes a cluster of swollen roots). Leaves opposite, sessile, or subsessile. Inflorescence umbel-like or raceme-like. Pedicel slender, short. Calyx with 5 basal glands. Corolla

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campanulate to subrotate; lobes erect or spreading, valvate. Corona double, attached to staminal column, outer series of 5 deeply 2-cleft lobes exceeding staminal column, inner series of 5 oblong lobes incumbent over stigma head, not or hardly exceeding staminal column. Filaments connate into a short tube; anthers without membranous apex; pollinia 2 per pollinarium, erect or ascending, with a translucent margin. Stigma head depressed. Follicles often linear, usually paired. Seeds comose.

Key to the species

1a. Erect herbs; not branching; flowers axillary or terminal umbels.......................2 1b. Climbing herbs; sometimes branching; flowers in axillary cymes..................29 2a. Corona without any processes, shortly toothed..................................................3 2b. Corona with five subulate lobes, undulate or toothed........................................7 3a. Flowers solitary; corolla deeply 5-lobed............................................................4 3b. Flowers 3-5.........................................................................................................6 4a. Flowers c. 2 cm in diameter; leaves c. 5 cm long........................B. attenuatum 4b. Flowers c. 0.5 cm in diameter; leaves c. 10 cm long..........................................5 5a. Corolla dark purplish, glabrous, margins recurved; corona 5-toothed������������������������������������������������������������������������ B. pauciflorum 5b. Corolla purplish brown, villous within, margin not recurved; corona irregularly toothed..........................................................B. parviflorum 6a. Corolla lobes narrowly lanceolate; cup-like outer corona raised above the corolla tube................................................................................... B. saldanhae 6b. Corolla lobes linear; corona lobes not raised above the corolla tube..................................................................................... B. glabrum 7a. Plants less than 5 cm high; corolla lobes more than 1.5 cm long, cage ­globular............................................................. B. ananthapuramense 7b. Plants more than 5 cm high; corolla lobes less than 1.2 cm long, cage conical................................................................................................... 8 8a. Corolla glabrous; stems pubescent at maturity...................................................9 8b. Corolla puberulous within; stems glabrous at maturity...................................10 9a. Corolla purplish brown; stamens and anthers deep ­yellow............. B. swarupa 9b. Corolla purple with yellow or white spots; stamens and anthers orange red: 10a. Leaves linear or linear-lanceolate; flowers in terminal umbels; corolla reflexed; outer corona cupular..........................................................................11 10b. Leaves ovate-oblong; flowers axillary, in 3-flowered cymes; corona 5-lobed, yellow................................................................................................................12 11a. Flowers solitary on axils; interstaminal corona lobes deeply divided, erect; pollinarium with a winged corpusculum..........................................B. vemanae 11b. Flowers 2-4 on axils; interstaminal corona lobes not divided, pendulous; pollinarium without winged corpusculum........................................B. ciliatum 12a. Leaves minute, subulate; flowers sessile......................................B. laevigatum 12b. Leaves filiform to lanceolate, acute..................................................................13 13a. Herbs up to 30 cm high; leaves to 2.8 cm long; flowers brightly ­colored........15

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13b. Herbs up to 80 cm high; leaves to 19 cm long; flowers pale white with black spots.............................................................................. B. nallamalayana 14a. Tubers small, 4 cm in diameter; flowers without purple spots on the corolla lobes..................................................................................................................14 15a. Outer corona 5-lobed, densely ciliate..........................................B. annamacharyae 15b. Outer corona cupular, slightly 5-lobed with sparse long white hairs.................................................................................................... B. naorojii 16a. Corolla lobes tapering; connate or twisted at the apex....................................16 16b. Corolla lobes free at apex; lobes not broad at base......................................... 24 17a. Plants up to 15 cm high; leaves up to 1 cm broad; flowers 1-3 in axillary node............................................................................................................. 17 17b. Plants up to 30 cm high; leaves up to 2.5 cm broad; flowers usually four.................................................................................................B. malwanense 18a. Peduncle present; cyme up to 5-flowered; corolla base urceolate, lobes free........................................................................................... B. rapinatianum 18b. Peduncle absent; cyme 3-flowered; corolla base campanulate, lobes free or connate.........................................................................................................18 19a. Corolla glabrous................................................................................................19 19b. Corolla hairy.....................................................................................................20 20a. Flowers pendulous; corolla connate, hairy at margins.............B. gondwanense 20b. Flowers erect; corolla spreading, densely hairy on the surface and ­margins..................................................................................... ..B. vartakii 21a. Corona glabrous................................................................................................21 21b. Corona hairy.....................................................................................................22 22a. Inflorescence 1–8-flowered, drooping; bract green, hairy at tip, p­ ersistent; sepals subulate; corolla lobes ovate, yellow but green at base; corona ­greenish yellow, blotched purple inside and o­ utside.................... B. mahajanii 22b. Inflorescence 2-flowered at every node, upright; bract greenish pink, ­glabrous, caducous; sepals triangular-acuminate; corolla lobes ­oblong-ovate, yellow, mottled dull/bright pink at base; coronal cup blackish ­yellow............................................................................ B. mahendragiriensis 23a. Thin-stemmed; staminal corona short; petals connate..................B. shrirangii 23b. Thick-stemmed; staminal corona long-lobed, deeply indented toward the ­center; petals spreading..............................................................................23 24a. Corolla deep yellow, slightly pubescent within; corona lobes notched above.................................................................................. B. megamalayanum 24b. Corolla purple with yellow tips, glabrous; coronal lobes simple................................................................................ B. matthewianum 25a. Flowers sessile or subsessile; corolla lobes linear............................................25 25b. Flowers peduncled; corolla lobes not linear.....................................................28 26a. Flowers 3–5 or less; corolla lobes 3 mm long, with purple hairs within.........26

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26b. Flowers 5 or more; corolla lobes 9–10 mm long, white villous within...........................................................................................B. rangacharii 27a. Corolla tube ash-colored, corona pure yellow................................ B. pullaiahii 27b. Corolla tube other than ash-colored, corona greenish or reddish yellow.........27 28a. Corolla tube with radial compressions, dark green colored, lobes with white hairs along the margins; corona cupular, staminal corona lobes shorter than anthers.................................................................................B. penchalakonense 28b. Corolla tube without radial compressions, green and red spotted, lobes with purple hairs along margins; corona annulate, staminal corona lobes projecting above anthers............................................................................... B. maculatum 29a. Corolla lobes narrowly triangular, hairs on lobes...........................B. bourneae 29b. Corolla lobes lanceolate, glabrous within.................................. B. elenaduense 30a. Corolla tube base quite flat, bulging limitedly with a constricted neck; more than 1 cm long..................................................................................................30 30b. Corolla tube campanulate; less than 5 mm long..............................................31 31a. Calyx segments 9–10 mm long; corolla tube ­urceolate; corona reddish maroon; follicle connate ­apically................................................ B. nigidianum 31b. Calyx segments 5 mm long; corolla tube slightly bulged; corona purplish; ­follicle divergent................................................................................ B. volubile 32a. Corolla tube glabrous; lobes connate at apex, glabrous, ­interstaminal corona glabrous; staminal corona flesh ­colored...................B. brevitubulatum 32b. Corolla tube with a ring of hairs around the corona and pubescent inside; lobes spreading, hairy; intrastaminal corona with hairs; staminal corona lemon yellow.......................................................................B. seshachalamense Brachystelma ananthapuramense Prasad et al., Kew Bull. 73: 16. 2018. Type: India, Andhra Pradesh, Ananthapuramu district, Gorantla hills, 800  m, 8 June 2014, A.N. Swamy & K. Prasad 44922 (holotype CAL; isotype SKU). Etymology: The new species is named after the type locality Ananthapuramu, which is a part of the Southern Deccan plateau of Andhra Pradesh. Dwarf perennial herb, 2–5 cm tall. Tubers globose or subglobose, c. 3 × 2 cm, pale white, with a few short roots. Stem short, erect, solitary, unbranched, pubescent, or glabrous; internodes c. 3 mm, short. Leaves light greenish brown, oppositedecussate, subsessile, oblong-lanceolate, ­elliptic-lanceolate, or ovate-lanceolate, 2–5 × 0.5–1 cm, apex acute or acuminate, margins and midrib below with hairs or glabrous. Inflorescence extra-axillary, sessile, solitary flower either side of the node; bract small, greenish pink, subulate, 0.2–0.4 mm long, acute, persistent, bracteoles absent. Flowers large, erect; pedicels short, greenish pink, cylindrical, 3–4 mm long, less than 1 mm broad, pubescent. Calyx gamosepalous, green or greenish pink, glabrous; sepals ovate-lanceolate or linear-lanceolate or subulate, 1–1.5 × 0.6–0.8 mm, acute. Corolla divided near to base, uniformly purple but darker inside; corolla tube much shorter than the lobes, c. 1 mm long; corolla lobes erect, connate at apex to form a globose cage, linear, 1.5–2.5  × 0.1 cm, acute, margins recurved, glabrous outside, and puberulous and sparingly white pilose within. Corona biseriate, 5-angled, uniformly dark purple, and densely

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pubescent; interstaminal corona cup-shaped, 2–2.5  mm across, 5-lobed near to the middle; lobes erect, triangular, thickened in middle, c. 1 mm long; staminal corona included in the interstaminal corona cup, 5-lobed; lobes  incurved, narrowly triangular, 0.6–0.8 mm long, incumbent, and adpressed on the anther lobes. Pollinium yellow, subglobose or oblong-ovoid, 0.2–0.4 × 0.2 mm, with pellucid lateral margin, basally attached to corpusculum by short, tubular caudicles. Follicles solitary or paired, cylindrical, tapering toward the apex, 6–8 cm long. Seeds comose, narrowly elliptic, 4–5 × 2–3 mm (Figures 2.1 and 2.2).

FIGURE 2.1  Brachystelma ananthapuramense Prasad et al.: (A) Habit, (B) Flower bud, (C) Calyx lobe, (D) Flower, (E) Flower without hairs, (F) Corona top view, (G) Corona side view, (H) Pollinia, and (I) Gynostegium. (Reproduced from Prasad, K. et al., Kew Bull., 73, 16, 2018. With permission from Trustees of the Royal Botanic Gardens, Kew.)

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FIGURE 2.2  Brachystelma ananthapuramense Prasad et  al.: (A) Habit, (B) Flowering plant, (C) Flower bud, (D) Flowers, (E) Tuber, (F & G) Calyx lobes, (H–K) Details of corona, (L) Pollinia, (M) Gynostegium, (N) Follicles, and (P) Seeds. (Reproduced from Prasad, K. et al., Kew Bull., 73, 16, 2018. With permission from Trustees of the Royal Botanic Gardens, Kew.)

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Found growing on black and red mixed soils in open, dry deciduous forests at an altitude of 800 m a.s.l. The common associates include Heteropogon contortus (L.) P. Beauv. ex Roem. & Schult., Tripogon bromoides Roth, Randia dumetorum (Retz.) Lam., Euphorbia fusiformis Buch.-Ham. ex D. Don and Drimia spp. Fl. & Fr.: May–July. Andhra Pradesh: Ananthapuramu distr., Gorantla hills, 800  m, A.N. Swamy  & K. Prasad 44965 (SKU), B.R.P. Rao & A.N. Swamy 45011 (SKU). Note: B. ananthapuramense is morphologically similar to the B. kolarense complex but differs by its small habit, short internodes, large flowers with more than 1.5 cm long corolla lobes, globular cage, and densely pubescent corona. IUCN Category: The species has a restricted distribution with more than 100 individuals in the Gorantla hills of Ananthapuramu, and further explorations in the adjacent hill tracts are necessary to ascertain its status. The current known habitat is prone to annual forest fires and grazing. Being edible, its tubers are harvested heavily by local people and also ploughed out by wild animals. It is currently categorized as Data Deficient (DD) (Prasad et al., 2018). Brachystelma annamacharyae Prasad et al., Nordic J. Bot. 34: 360. 2016. Type: Andhra Pradesh, Kadapa Distr., Sanipaya, P.V. Prasanna, K. Prasad, M. Sankara Rao and T. Thulasaiah 4568 (Holotype BSID, Isotype BSID). Etymology: The new species is named after the renowned saint ­composer Sri Tallapaka Annamacharya, who has composed devotional songs in praise of Lord Balaji, the presiding deity of Seshachalam hills. Dwarf, perennial herb, 4–12 cm tall. Tubers globose, c. 4 × 4 cm, dirty brown. Stem short, erect, solitary, unbranched, 1.5–2 mm thick, in juvenile stage pubescent becoming glabrous at maturity; internodes c. 1.5 cm long. Leaves light greenish brown, simple, opposite-decussate, petiolate; petiole 1–2 mm long, glabrous; lamina lanceolate, ovate-lanceolate, ovate-oblong, 2–5.5 × 1–2.2 cm, acute, with puberulent margins; midrib puberulent and prominent below. Inflorescence extra-axillary, sessile, solitary; bract one, pinkish, linear, 0.6–0.8 × 0.2 mm, acute, glabrous, caducous. Flowers horizontally spreading with long pedicel. Pedicel cylindrical, filiform, 3–3.5  cm long, pubescent. Calyx gamosepalous, glabrous, greenish yellow, lobes ovate, 1 × 0.5 mm, acute or obtuse. Corolla yellowish, spotted with purple, tube short, campanulate, pilose, c. 1.5 mm long; corolla lobes ovate, 2–2.5 × 1 mm, acute, reflexed, margins slightly recurved, densely pilose inside and glabrous outside; trichomes pinkish white, 2–3 mm long. Corona biseriate, staminal and interstaminal parts fused to a cup-shaped structure, c. 2 mm across, basally bright yellowish; interstaminal corona lobes erect, purplish, much taller than gynostegium, oblong, keeled, 0.8 ×  0.4 mm, rounded at apex, with few white trichomes at apex margins; staminal corona basally fused to interstaminal corona ring, its lobes triangular, c. 0.4 mm long. Pollinium subglobose, c. 0.2 × 0.18 mm, is yellow, with pellucid lateral margin, basally attached to c. 0.1 mm long corpusculum by light-brown, short, tubular caudicles. Follicles solitary, purplishyellow, cylindrical, tapering towards apex, 8–9 cm long; fruiting pedicels straight, spreading to erect. Seeds comose, narrowly elliptic, 4–5 × 1.5–2 mm, light brown at center (Figures 2.3 and 2.4). Endemic to the Seshachalam hills in the Kadapa district of Andhra Pradesh. Fl.: June–September; Fr.: December–January

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FIGURE 2.3  Brachystelma annamacharyae Prasad et al.: (A) Habit, (B) Flowering twig, (C) Calyx lobe, (D) Flower, (E) Flower without hairs, (F) Pollinia, and (G) Seed. (After Prasad, K. et al., Nordic J. Bot., 34, 360, 2016. With permission.)

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FIGURE 2.4  Brachystelma annamacharyae Prasad et al.: (A), (B), Habit in August–September, (C) Habit in December, (D) Flower, (E) Top view of flower, (F) Side view of column, (G) Top view of column, (H) Top view of gynostegium, and (I) Pollinia. (After Prasad, K. et al., Nordic J. Bot., 34, 360, 2016. With permission.)

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India: Andhra Pradesh. Endemic. IUCN Category: Following the IUCN guidelines ver. 11 (IUCN, 2014), Brachystelma annamacharyae is assessed as “Data Deficient” (DD), as only a few individuals were located during the floristic survey. Further explorations in the adjacent hill tracts are necessary to ascertain the status of the species. Presently, the area is well protected as it falls under the Seshachalam biosphere reserve. Therefore, there is no immediate threat to this species, but the habitat is prone to seasonal, anthropogenic forest fires and to grazing (Prasad et al., 2016). Affinities: Brachystelma annamacharyae is most closely allied to B.  ­s warupa, as is expressed by the possession of small flowers with reflexed corolla lobes and an exposed corona. Brachystelma annamacharyae differs from B. swarupa in certain characters as tabulated (Table 2.2). This species is easily distinguishable by the dense indumentum of the adaxial corolla surface of long pinkish white hairs, a character that is very rare in the genus as a whole. Another Indian species, Brachystelma ciliatum Arekal & T.M. Ramakrishna (1981) from Karnataka and Andhra Pradesh, is obviously also related to these two species, but it possesses 2–4-flowered cymes, ciliate corolla lobe tips, and a short, flattened corona. Vegetatively, B. annamacharyae is also unusual with its dwarf growth; the single stem reaches only a few centimeters in height and the light greenish brown colored leaves are more or less ovate (vs. green, narrowly lanceolate in

TABLE 2.2 Morphological Comparison of Brachystelma annamacharyae and B. swarupa Characters

B. swarupa

B. annamacharyae

Plant height (cm) Leaves (cm)

30–60 Green, subsessile, narrowly lanceolate, 6–12 × 0.3–0.7

Inflorescence Calyx lobes Corolla tube

24–28-flowered Puberulent on the outside Less than 1 mm long, flat to reflexed

Corolla lobes

Elliptic-ovate, glabrous, 4–6 mm long cilia present near the rim of the tube, completely reflexed when mature Tip bifid, glabrous, margins long-ciliate Without free tips

4–12 Greenish brown, petiolate, lanceolate, ovate-lanceolate or ovate-oblong, 2–5.5 × 1–2.2 1-flowered Glabrous 1.5 mm long, campanulate, not reflexed Ovate, adaxially densely covered with purplish white trichomes, more or less reflexed Apex entire, obtuse, margins with few long trichomes With free tips

Laterally with wing-like projections

Without projections

Interstaminal corona Staminal corona lobes Corpusculum

Source: Prasad, K. et al., Nordic J. Bot., 34, 360, 2016.

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B. swarupa). Habitually and florally, however, the two South African species— Brachystelma arnottii Bak. and B. minimum R.A. Dyer—show most congruence with B. annamacharyae (cf. Illustrations in Dyer, 1883). Both represent dwarf herbs only a few cm high, with small flowers possessing recurved corolla lobes. In particular, B. minimum resembles B.  annmacharyae with its single-flowered inflorescences bearing the little flower on a long stalk. But there, as in B. swarupa as well, the corolla is nearly tubeless and the corona exposed, and corolla surfaces are glabrous. However, the considerable morphological similarities between these species are obviously due to parallel evolution and not an expression of phylogenetic closeness (Prasad et al., 2016). Brachystelma attenuatum (Wight) Hook. f., Fl. Brit. India 4: 65. 1883; Albers & Meve, Ill. Handb. Succ. Pl. Asclepiadaceae 21. 2002. Eriopetalum attenuatum Wight, Contr. Bot. India 35. 1834. Type: s.coll. s.n. (K000794639) Etymology: attenuatum = tapering, narrowing; in allusion to corolla segments being filiform and 4–5 times longer than the corolla tube. Segments of corolla filiform, 4–5 times as long as the tube, villous, corona shortly irregularly toothed. Flower 2 cm in diameter. North-Western India. Native of West Himalayas. Note: J.D. Hooker (loc. cit.) mentioned that he has seen only a single flower in Herb. Wight. IUCN Category: This species has not been recollected after the type collection. It is probably extinct. Brachystelma bourneae Gamble, Bull. Misc. Inform. Kew 1922: 120. 1922 & Fl. Madras 2: 851. 1923; S.R. Srinivasan in Henry et al., Fl. Tamil Nadu Ind., Ser I: Analysis 2: 80. 1987; A.P. Jagtap & N.P. Singh, Fasc. Fl. India 24: 179. 1999. Type: India: South India: Madurai Distr., Pulney hills, on Kodaikanal Ghat and in Perumal and Vailpatti Valley, east slope, May 1898 and June 1899, Sir A.G. and Lady Bourne 1020, 2751, 2752 (K) Etymology: The name honors Lady Emily Tree Bourne; she was a well-known botanical artist. Her illustrations were used in the “Flora of the Nilgiri and Pulney Hill-tops” by P.F. Fyson. Herbs erect, c. 30 cm tall, tuberous. Tubers fusiform, 3–4 cm in diam. Leaves linear, 7–10 × 0.2 cm, opposite, glabrous above and puberulous below, midrib prominent, lateral veins obscure, margin puberulate, reddish; petiole subsessile or 0.1 mm long, puberulate. Flowers in cymes, axillary, sessile, 3-flowered. Pedicels filiform, 5–15 mm long, glabrous. Bracteoles minute. Calyx 5-lobed, 0.1 × 0.4 mm, lobes linear, glabrous, acute, 2 mm long. Corolla tube short, rotate, 0.2 mm long, white with dark green spots, lobes triangular-lanceolate, 0.1 × 0.3 mm, acute, purple hairs on the lobes and margins, corona purple, lobes very thin, ciliate, torti apex, c. 1 cm long, inflexed. Stamens obtuse, pollinia globose, c. 0.5 × 1.2 mm, yellowish with pellucid margins. Style apex pentagonal (Figure 2.5).

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FIGURE 2.5  Brachystelma bourneae Gamble: (a) Habit, (b) Flower—front view, (c) Flower-lateral view, and (d) Tuber. (Courtesy of Karuppusamy.)

Rare in Dry deciduous forests of the Western Ghats (Eastern slopes of Palni hills on Kodaikanal Ghat and in Perumal and Vilpatti valley) in Tamil Nadu. Fl. & Fr.: May–June Tamil Nadu: Dindigul Distr., Kodaikanal Taluk, Perumalmalai, Grass peak, S. Karuppusamy 1244 (SGH). India: Tamil Nadu. Endemic. IUCN Category: The fresh collection of this species is not available at MH or CAL. The second author collected this specimen recently in the Perumal Malai Peak of Palni hills, which showed the rediscovery of this species after a gap of 120 years from the type locality.

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Brachystelma brevitubulatum (Bedd.) Gamble, Fl. Madras 2: 852. 1923; S.R. Srinivasan in Henry et al., Fl. Tamil Nadu Ind., Ser I: Analysis 2: 80. 1987; A.P. Jagtap & N.P. Singh, Fasc. Fl. India 24: 179. 1999; Albers & Meve, Ill. Handb. Succ. Pl. Asclepiadaceae 23. 2002; Vijayasankar et al., Phytotaxonomy 3: 130. 2003; Pullaiah et  al., Fl. Eastern Ghats 4: 91. 2011. Ceropegia brevitubulata Bedd., Ic. Pl. Ind. Or. t. 174. 1839. Brachystelma beddomei Hook.f., Fl. Brit. India 4: 66. 1883. Type: Tamil Nadu: Amerdy hills, near Vellore, 1000 ft., Beddome 232 (flowers & fruits), December 1871 (K). Etymology: Short-tubed Brachystelma (short corolla-tubed Brachystelma). A weak-stemmed, tuberous, annual, twining herb. Sap colorless, watery. Tubers depressed-globose, 1.5–2 × 2.5–3 cm, dull brown outside, whitish inside, with transparent thin skin. Stems unbranched, slender, cylindrical, twining, internodes 7–10 cm long. Leaves opposite, pairs often unequal, linear-lanceolate, 8–11 × 0.4–0.8 cm, thinly chartaceous, puberulous above, glabrous beneath, with scattered hairs on nerves beneath along with s­ cattered white dots, cuneate and with a brownish gland at base, apex gradually a­ cuminate, margin minutely ciliate and slightly thickened along margin, midrib grooved above, raised beneath, reticulations distinct; lateral nerves indistinct, alternate, 9–14 pairs, joining along margins; petioles 3–8 mm long, hyaline, ciliate, dilated at base. Stipules axillary, 3-fid, setaceous, up to 1 mm long. Inflorescence axillary and terminal, umbellate cymes, 3(4)-flowered; peduncles 2–3 mm long, puberulous, pedicels 5–10 mm long, hyaline,  puberulous. Bracts 2 , setaceous, c. 3 mm long, puberulous. Flowers c. 3.5 cm long and c. 1.5 cm across. Calyx 5-lobed, free from base, lobes linear-subulate to lanceolate, long acuminate at apex, 2–5 mm long, hyaline along margins, glabrous within, scattered hairy on nerves without. Corolla tube short, connected at base and apex, up to 3 mm long, lobes 5, ovate-lanceolate, 2–3.5 × 0.5–0.8  cm, subfleshy, long-acuminate at apex, cohering at their tips, purplebrown within, greenish yellow without. Coronal processes entirely confluent with anthers. Follicles usually paired, almost equal, narrowly cylindrical, c. 7.5 × 0.4 cm, gradually tapering towards apex, blunt at tip, greenish with purple tinge, with thin powdery coating. Seeds c. 10 in each follicle, oblong, compressed, 0.5 × 0.2 cm, brownish, coma silky white, terminal, unequal, up to 3 cm long, smooth (Figure 2.6). Rare in dry deciduous forests. Fl. & Fr.: November–January. Tamil Nadu: Tiruvannamalai Distr., Polur Taluk, Paruvathamalai R.F., R.  Vijayasankar 71248 (flowers), 10-11-2002 (FRLH), R. Vijayasankar  & K. Ravikumar 71358 (fruits, 12-01-2003 (FRLH); Chengam taluk, Swamimalai R.F., R. Vijayasankar 59183 (flowers), 10-10-2003 (FRLH). India: Tamil Nadu. Endemic. Note: Beddome first collected this species during 1871 from the Thellay hills of Amerdy (Amirthi) forest near Vellore at about 350 m altitude; the type specimens are housed in the Kew Herbarium. The Amirthi forest, though close to Vellore, falls in the Tiruvannamalai district in Tamil Nadu. To realize the beauty and rarity of this species, it is imperative to quote Beddome’s words from his protologue “…this beautiful rather abnormal species appears to be very rare” and he had seen only one plant. He further adds that his attempt to grow the plant by means of tuberous roots at his

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FIGURE 2.6  Brachystelma brevitubulatum (Bedd.) Gamble: (A) Habit, (B) Flower bud, and (C–F) Flowers in different angles. (Courtesy of Vijayasankar Raman.)

residence were futile. Vijayasankar et al. (2003) collected this species in 2002, from the Tiruvannamalai district nearly 131 years after the type collection from a locality other than the type locality. Ramakrishna (1995) recorded this species in Karnataka and this report is based on Hooker’s Flora of British India, which is once again based on Beddome and states “Mysore; Vellore hills alt. 1000 ft, Beddome.” Vellore hills, as given above, is in Tamil Nadu and not in Karnataka. Hence, report of this species from Karataka is not correct; it does not occur in Karnataka. Perhaps based on this report, P. Singh et  al. (2015) record this species also from Karnataka, but we could not locate any Herbarium specimen in any Indian Herbaria and have not recorded this from Karnataka.

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Nomenclatural Note: The Plant List database lists the name Brachystelma beddomei as the accepted name based on the reference Fl. Brit. India 4: 66 1883 http://www.theplantlist.org/tpl1.1/record/kew-2681511. This data has been derived from the World Checklist of Selected Plant Families (WCSP), which includes the name Ceropegia brevitubulata Bedd. as an accepted name. As per IPNI, the name B. beddomei Hook.f. is illegitimate and superflous. Therefore, B. brevitubulatum (Bedd.) Gamble should be the correct accepted name. IUCN Category: Not assessed. Brachystelma ciliatum Arekal & T.M. Ramakrishna, Curr. Sci. 50: 145. 1981; T.M. Ramakrishna et  al., Asclepiadaceae Karnataka 38. 1995; Albers & Meve, Ill. Handb. Succ. Pl. (Asclepiadaceae) 2: 30. 2002; Kullayiswamy et al., Nelumbo 55: 191. 2013 & Fl. Thummalapalli Uranium Mining Area 256. 2017. Type: Karnataka, Kolar Distr., Sonnipally 920  m, 27 June 1979, Ramakrishna 1311 A (Holotype CAL, Isotype BSI!, K); Ramakrishna & Arekal 1340 A-E (Paratype, BSI!, Mysore Univ.); Ramakrishna 1316 (Paratype, BSI!). Etymology: Brachystelma ciliatum = ciliated Brachystelma; in allusion to corolla lobe tips being ciliated. Vern.: Telugu: Wootakunimmata gadda, Pithu Jougu Nimmatayalu Perennial tuberous herb. Tuber brown, subglobose–globose, 4 cm across and dorsiventrally compressed, 5.6 × 5.6 × 1.5 cm. Stem erect, 30–35 cm high, 1.5–2 mm in dia., rarely branched, purplish, pubescent, buried part of the stem 3–4 cm, white, glabrous. Leaf scales opposite, 3–4 pairs from the base, ovate, c. 3 × 2  mm, brown puberulent, sessile; leaves opposite in middle of the shoot, 4–5 pairs, linear, linear–lanceolate, 5–9 × 0.2–0.3 cm, nerves 9–12 pairs when young, semi-succulent, brown, midrib pubescent beneath, base acute, margins entire, apex acuminate, pubescent, sessile with 2–3 ovate glands at base, greenish pink; epidermal hairs unicellular. Inflorescence axillary, 2–4-flowered umbel, pendulous. Pedicels filiform, 3.38–3.81 cm long, pubescent, stout and erect in fruits; bracts 4 at each node, each c. 1  mm, ovate, pubescent. Calyx 5-lobed, minute, c. 0.5 mm, connate at base, ovate, pubescent. Corolla 5-lobed, connate at base, lobes oblong to ovate, c. 5 × 2 mm, glabrous, narrowing at apex with 2 mm long shiny cilia, greenish yellow, dark pink at apex and middle; tube c. 1 mm long, lobes reflexed when mature. Corona 2whorled; outer whorl 5-lobed, c. 2.5 mm across, cupular, glabrous, dark pink dotted, 10-lobed at margin; inner whorl 5-lobed, alternate with stamens, ovate–oblong, extended to staminal c­olumn. Staminal column 5-lobed, yellow, elliptic, winged; stigma light greenish yellow, pentagonal, truncate, minute, sticky. Pollinarium yellow, 60 µm across, erect; corpusculum brown. Follicles 1–2, linear-lanceolate, 7–8 × 0.2–0.3 cm, light reddish brown, glabrous, unequal, curved at apex when young, acuminate and stronger when mature. Seeds 8–12, brown, ovate, 7–8 × 2–25 mm, thin; coma white, 2–2.5 cm long (Figures 2.7 and 2.8). Rare in the Anantapur district, Andhra Pradesh associated with Ophioglossum gramineum, Heteropogon sp., Chrysopogon sp., etc., in sandy moist places of dry deciduous forests. In Karnataka, it occurs in a 7  km area in the Kolar district in

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scrub jungle usually associated with short grasses and among dicotyledons such as Canthium parviflorum and Dodonaea viscosa. Fl. & Fr.: August–October Chromosome No: 2n = 22 Andhra Pradesh: Anantapur Distr., Nigidi forest, KRKS 41870  A—D (SKU, MH & CAL); Batrepalli, S.S. Kambale & K. Raja Kullayi Swamy 250 (SUK);

FIGURE 2.7  Brachystelma ciliatum Arekal & T.M. Ramakrishna: (A) Twig, (B) Flower, (C) Corolla dorsal view, (D) Corolla ventral view, (E) Corona, and (F) Young fruit. (Courtesy of K. Raja Kullayiswamy.)

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FIGURE 2.8  Brachystelma ciliatum: (A) Habit, (B) Flowers, (C) Habit with association, (D & E) Foliar glands, (F) Young fruit, (G) Mature fruit, (H) Persistent fruit layers, (I & J) Seeds, (K) Deltoid scales on shoot base, and (L & M) Tubers. (Courtesy of K. Raja Kullayiswamy.)

Kalasamudram, Kadiri, S.S. Kambale & K. Raja Kullayi Swamy 99 (SUK). Karnataka: Kolar Distr., Sonnipally, T.M.  Ramakrishna 1178, 1221, 1311, 1312, 1314, 1315, 1316, 1332, 1341 (Mysore Univ.), T.M. Ramakrishna & D.A. Govindappa 1340 (Mysore Univ.). India: Andhra Pradesh, Karnataka. Endemic. Conservation status: As per IUCN Red list (IUCN, 2001); it is Vulnerable VU, D 1 + 2. The extent of occurrence of the species is estimated to be less than 3 km2

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in Andhra Pradesh and 7 km2 in Karnataka, that too, in a severely fragmented and degraded habitat with less than one thousand individuals. The tubers are a favored food of several wild animals and this pressure poses a real threat to the species (Kullayiswamy et al., 2013). Note: Plants of Brachystelma ciliatum are growing among grasses and are very difficult to locate in the vegetative and immature fruit stage. The plant can be recognized easily in the field in flower and in fruit, particularly in dehiscence. Brachystelma elenaduense Char, Proc. Indian Sci. Congr. Assoc. 58: 435. 1971 ex Satyan. Curr. Sci. 47: 965. 1978 (as  “elenaduensis”); G.D. Arekal & T.M. Ramakrishna, Bull. Bot. Surv. India 24: 217, 1982; Kartikeyan & Sharma, J. Bombay Nat. Hist. Soc. 80: 70. 1983; T.M. Ramakrishna et al., Asclepiadaceae Karnataka 40.1995; A.P. Jagtap & N.P. Singh, Fasc. Fl. India 24: 183. 1999; Kaliamoorthy et al., J. Threat, Taxa 7: 7587. 2015. Type: India: Karnataka, Tumkur Distr., Sandy grass at Elenadu, alt. 780.33  m, M.B.S. Char 1 (Holotype, CAL, Isotype University of Mysore Herbarium). Etymology: The specific epithet is derived from the place of the type collection, Elenadu in Karnataka state. Vern.: Kannada: Unnikana, Ullikana gadde Erect perennial herb, with tuber-like rhizome. Tubers ovate to discoid, brownish fleshy, 2–4 cm in diam., often compressed, upright in the middle. Stems 4.5–11 cm high, sparingly branched, slender, puberulous, hairs multicellular, uniseriate, internodes c. 0.4–0.65 cm long; buried part of the stem smooth and whitish in color. Leaves 1–2.5 cm × 0.6–3 mm, fleshy, linear to linear-lanceolate, apex narrowly acute, subsessile, glabrous above, puberulous beneath, and ciliolate along the margins. Flowers usually solitary, axillary, 5–9 mm long, greenish outside, velvety purplish within. Pedicel slender, 0.4 cm long, puberulous. Calyx 5-lobed, lobes 3–4 mm long, narrowly acute. Corolla rotate, purple, deeply 5-fid, reflexed, lobes 4–6 mm long, deltoid, spreading and reflexed at bloom, lanceolate, purple. Corona biseriate, outer corona cupular, 1–2 × 2–3 mm, obtusely 5-angled around the staminal corona and as high, undulate, purplish and with white hairs outside, teeth 5, subulate, each adnate to the base of corresponding staminal coronal scale; inner corona 5-lobed, tridentate, overtopping the anthers, cream colored, hairy. Pollinia 0.03 mm long, globular-oblong, inner-face winged, corpusculum brownish, caudicle tubular, with membranous structure on either side of corpusculum. Carpels free, glabrous, 2 mm long, stigma pentangular. Follicles 2, rarely 1 per flower, 4–8 cm long, slender. Seeds 8–10 × 3.5 mm, compressed, with an apical tuft of hairs, 2–3 cm long (Figure 2.9). Rare, growing along with short grasses in sandy soil in Elenadu in the Tumkur District of Karnataka. It was also found growing amidst grasses on open rocky hillslopes deposited with sandy loamy soil at an altitudinal range between 1200–1350 m in the Kolli Hills and Yercaud Hills of Tamil Nadu. The associated grass species were invariably found to be a Cymbopogon species. A scanty population of mature individuals was observed in the study areas. Karnataka: Tumkur Distr., Korekere, T.M.Ramakrishna 1338, 1339 (Mysore Univ. Herbarium). Tamil Nadu: Kolli Hills, Namakkal Distr., ~1200 m coll. Kaliamoorthy et  al. (National Orchidarium, Yercaud); Shevaroy Hills, Salem District, ~1350  m, coll. Kaliamoorthy et al. (National Orchidarium, Yercaud).

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FIGURE 2.9  Brachystelma elenaduense Sathyan: (A) Fruiting plant, (B) Flower, (C) Calyx, (D) Staminal corona, (E) Carpels, (F) Stigma top view, (G) Anther with inner corona, (H) Pollinia, (I) Fruit, and (J) Seed. (After Ramakrishna, T.M. et al., The Asclepiadaceae and Periplocaceae of Karnataka, Prasaranga, University of Mysore, Mysore, India, 1995. With permission.)

Fl.: April–June; Fr.: May–September Chromosome No. 2 n = 22 India: Karnataka, Tamil Nadu. Endemic. Note: In the original description, the author has described the outer series of staminal corona as petaline corona. This feature is not present in the genus Brachystelma. IUCN Category: Rare. Not assessed.

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Brachystelma glabrum Hook. f., Fl. Brit. India 4: 65. 1883; Gamble, Fl. Madras 2: 851. 1923; A.P. Jagtap & N.P.Singh, Fasc. Fl. India 24: 183. 1999; Albers & Meve, Ill. Handb. Succ. Pl. Asclepiadaceae 30. 2002; Pullaiah et al., Fl. Eastern Ghats 4: 2011. Type: India, Andhra Pradesh, Cuddapah (now called Kadapa) hills, Beddome s.n. (K000820004). Etymology: Brachystelma glabrum = glabrous or smooth Brachystelma; in allusion to the whole plant being almost glabrous. A short slender herb, glabrous, unbranched, to 30 cm high, minutely pubescent, latex watery. Leaves opposite, decussate, linear-lanceolate, 1–2.8 × 0.15–0.2 cm, minutely pubescent, apex and base acute, margin ciliate, lateral veins inconspicuous, petiole 0.5 mm long. Inflorescence terminal, few (3–5)-flowered umbels, peduncle 1.5 mm long. Flowers bright orange red. Pedicels filiform, slender. Calyx 5-lobed, divided nearly up to the base, lobes c. 1.5 × 0.5 mm, linear-lanceolate, tip pointed, minutely pubescent without, glabrous within, margin ciliate. Corolla slightly reflexed, lobes 5, linear, glabrous, three times as long as tube. Corona staminal, uniseriate, cupular, without subulate processes, c. 1 mm high, shortly toothed, without subulate processes, glabrous. Stamens 5, c. 1.5 mm long. Pollinia 5, yellow, waxy, c. 144 × 86.4 µm, with pellucid layer attached by light brown caudicles. Follicles paired, to 4.5 cm long, linear, glabrous. Seeds oblong, dark brown with light brown margin, c. 0.5 mm wide, base rounded, coma silky whit, c. up to 2 cm long. Rare in dry rocky hills. Fl.: June–August; Fr.: July–September. Andhra Pradesh: Cuddapah hills, Beddome s.n.(MH); Tamil Nadu: Shevaroy hills (SM), E.Vajrvelu 82557 (MH), S.Revathi Lakshmi & T.Senthil Kumar s.n. (MH); Kakkashola, Shevaroy hills (SM), E.Vajravelu 77732, 77740 (MH). India: Andhra Pradesh, Kerala and Tamil Nadu. Endemic. IUCN Category: Rare, not assessed. Brachystelma gondwanense Govekar et al., Rheedea 26: 145. 2016. Type: India, Maharashtra, Gadchiroli district, Lohara village, R.S. Govekar 174 (Holotype CAL; Isotypes BAMU, BSI, MH, SUK). Etymology: The species is named after Gondwana, the area including the type locality, which was once dominated and ruled by the Gond tribe. Perennial herb, erect, to 25 cm high, Tuber generally globose, 3–5 cm, greyish white with scattered, few roots (during summer, tubers in fruiting shoot are comparatively much smaller and shaped differently ranging from horizontally ellipsoid to pyriform or reniform). Stem solitary, terete, 1–2 mm in diam., generally unbranched, ridged, glabrous, whitish at base, glaucous green above; internodes 2–3 cm long, gradually decreasing in length near apex. Leaves simple, sessile, opposite-decussate in 3–7 pairs, lanceolate, 10–13 × 1.5–2 cm, smooth at margins, acute at apex, glabrous, green above, glaucous pale beneath; midvein strong and thicker at base, thin towards tip, protruding on lower side, very yellowish white; secondary veins 4 or 5, obscure on upper surface. Flowers on a long leafless, glabrous, yellowish green shoot of 8–12 x c. 0.2 cm, which turns dark in fruiting stage. Inflorescence initially crowded at apical nodes with internodes increasing gradually 4–6 cm; lateral, shortly peduncled, pubescent, 2-flowered, and inverted; pedicels stout, to 7 mm, minutely pubescent. Bracts and bracteoles persistent, linear, acute

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at apex, minutely glandular, greenish yellow, turning dull brown in fruiting stage; bracts c. 8 × 0.5 mm; bracteoles c. 3 × 0.3 mm. Calyx 5-lobed; lobes triangular, 3–4 × 0.5–1.5 mm, united at base, acute at apex, glandular-hairy, yellowish green. Corolla sharply angled conical in bud stage; tube to 1 mm long, shallow, 5-lobed at apex; lobes erect but twisted while opening and open gradually from base to tip and tips often remain united for a few days, linear, subcylindrical, 14–18 × 1.5–2 mm, involute at margins, acute at apex, 2–3 mm long white hairs scattered throughout the inner surface and margins but not on the outer surface, more dense toward lower half and almost absent near tips; lobes pale yellow with purple irregular blotches and yellowish green above with few purplish blotches only towards base. Corona biseriate—staminal and interstaminal parts fused and form bowl-shaped staminal corona. Interstaminal corona c. 3 mm across, slightly above the gynostegium of 5 shallowly bifid lobes, hairy inside, glabrous otherwise, yellowish white with purple dots; staminal corona is incumbent on anther sacs more than half the length of sacs, lanceolate, white-blotched with deep purple. Pollinarium c. 0.25 × 0.15 mm; pollinium ovoid, attached to red corpusculum by short caudicles. Follicles in pairs, erect, crowded at top, 8–10 × 0.25–0.3 cm, acute at apex, broader towards basal half, pinkish, smooth. Found mostly in dry to moist deciduous, lowland mixed forests with clayey gravelly to loamy soils at an elevation of 160–180 m and annual rainfall about 1500 mm in Maharashtra. Fl. & Fr.: April–June. Maharashtra: Gadchiroli district, Compartment No. 316, Chaprala Sanctuary, near Dhannur village, M.M. Sardesai 2014 (BAMU). India: Maharashtra. Endemic. Note: The tubers look like a small potato and are uprooted, peeled off, and eaten by the local people. This is one of the major threats to this extremely rare species. IUCN Category: Extremely rare. Brachystelma kolarense Arekal & T.M. Ramakrishna, Proc. Indian Acad. Sci., Pl. Sci. 90: 203. 1981; T.N. Ramakrishna et  al., Asclepiadaceae Karnataka 42. 1995; A.P. Jagtap & N.P. Singh, Fasc. Fl. India 24: 184. 1999; Albers & Meve, Ill. Handb. Succ. Pl. Asclepiadaceae 33. 2002; Kambale et al., Nat. Acad. Sci. Lett. 40: 301. 2017. Type: India: Karnataka, Kolar Distr., Sonnipally, T.M. Ramakrishna 1323 (Holotype CAL; Isotype Mysore Univ.!). Etymology: The specific epithet is derived from type locality Kolar in the Karnataka state. Perennial, erect, tuberous herb, 8–15 cm high. Tubers flattened or ovate, fleshy, brownish, 2.5–4 cm across. Stems erect, sparingly pubescent. Leaves subsessile, ovate-oblong to ovate-lanceolate, 1.8–4.5 × 0.8–1.5 cm, apex acuminate, base cuneate, glabrous above, puberulous beneath, more so on the midrib. Inflorescence axillary, two-flowered cymes. Peduncles 4 mm long, puberulous. Flowers purple spotted outside and purplish black within. Bracts linear, 1–3 mm long, caducous. Pedicels 2–4 mm long, puberulous. Calyx 5-lobed, divided up to a third way down, pubescent outside, glabrous within. Corolla 5-lobed, lobes 4–5 mm long, narrow,

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acute, connate at apex, margin recurved at base so as to expose the corona, glabrous outside, puberulous within. Corona 2-seriate, staminal, stipitate, outer corona lobes 5, bifid, 0.5 mm long, hairy; inner corona arising on the inner face of outer corona, partially covering anthers. Anthers flattened, incumbent over the stigmatic head, apex truncate; pollinia erect, 270 µm long, ovate-oblong, winged on inner face; corpusculum dark brown. Carpels  free, oblong, stylar apex pentagonal, flat (Figures 2.10 and 2.11). Rare, growing in crevices of rocks. Chromosome No. 2 n = 22 Karnataka: Kolar Distr., Sonnipally, T.M.Ramakrishna 1310 (Mysore University Herbarum); Khanapur, S.S.Kambale SSK 344 (SUK). Kerala: Kasaragod district, Periya, S.S.Kambale, A.Gholave & A.A.Adsul SSK 349 (SUK). India: Karnataka and Kerala. Endemic. Note: The connate tips of corolla with recurved margins at the base recall the condition of the corolla of Ceropegia. IUCN Category: Rare. Brachystelma laevigatum (Wight) Hook.f., Fl. Brit. India 4: 65, 1883; Duthie, Fl. Upper Gang. Pl. 1: 512. 1960 (repr. ed.); T.M. Ramakrishna et al., Asclepiadaceae Karnataka 44. 1995; A.P. Jagtap & N.P. Singh, Fasc. Fl. India 24: 185. 1999; Albers & Meve, Ill. Handb. Succ. Pl. Asclepiadaceae 33. 2002. Eriopetalum laevigatum Wight, Contrib. Bot. India 35. 1834. Gomphocarpus laevigatus Buch.- Ham. ex Decne, in DC. Prodr. 8: 563. 1844. Type: Gomphocarpus laevigata Buch.- Ham. in Sylvis Cossale, Gorukpur, F. Hamilton s.n. (K000820017). Etymology: Brachystelma laevigatum: smooth/shining Brachystelma; laevigatum = smooth, free from wrinkles, hairs, etc., often shining; in allusion to the corolla being smooth. Erect, perennial, tuberous herb, tubers up to 2.5 cm across, flattened upright. Stem 30–45 cm high, unbranched, pubescent upwards; hairs multicellular, uniseriate. Leaves reduced to scales, scales linear, acicular, 2–5 mm long, brownish. Inflorescence axillary, sessile, 1–3-flowered umbels. Flowers pendulous, c. 1 cm in diam. Pedicels minute, hispid. Calyx 5-lobed, lobes divided up to two-thirds way down, ovate, acute, pubescent outside, glabrous within. Corolla reflexed, pale dull purple, with pale green, filiform segments three times as long as the tube, puberulous outside. Corona 2-seriate, staminal, cup-like, sessile; outer corona 5-fid, lobes up to 0.5 mm long, puberulous; inner corona 5-lobed, arising on the inner face of the outer corona forming second series, lobes 1 mm long, incumbent, and partially covering the anthers; anthers up to 1 mm long, incumbent over stigma, thecae lateral, apex inapediculatee, truncate; pollinia 275 µm long, ovate-oblong, winged within, corpusculum brownish, caudicle tubular; carpels free, oblong, glabrous, stigma pentangular, flat. Follicles 5–8 cm long, erect, subfalcate, tapering gradually to the apex. Seeds c. 0.5 cm long, linear, margined, coma twice as long as the seed (Figure 2.12). Rare, growing under the shade of Canthium parviflorum and Dodonaea viscosa in gravelly soil. Fl. & Fr.: March–April.

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FIGURE 2.10  Brachystelma kolarense Arekal & Ramakrishna: (A) Flowering plant, (B) Flower, (C) Calyx, (D) Staminal corona, (E) Carpels, (F) Stigma top view, (G) Anther with inner corona, and (H) Pollinia. (After Ramakrishna, T.M. et al., The Asclepiadaceae and Periplocaceae of Karnataka, Prasaranga, University of Mysore, Mysore, India, 1995. With permission.)

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FIGURE 2.11  (A) Brachystelma kolarense Arekal & Ramakrishna, (B) Brachystelma malwanense, (C) Brachystelma naoroji, and (D) Brachystelma shrirangii. (Courtesy of Sushant More with permission.)

Karnataka: T.M. Ramakrishna 1317 (Mysore University Herbarium). India: Gujarat, Karnataka and Uttar Pradesh (Gorakhpur District). Endemic. IUCN Category: Duthie (1960) described the only specimen collected by Buchanan-Hamilton in the sub-Himalayan districts of Gorakpur during 1814. Since then, there were no reports of its collection until T.M. Ramakrishna collected the same in 1995.

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FIGURE 2.12  Brachystelma laevigatum (Wight) Hook.f.: (A) Flowering twig, (B) Flower, (C) Flower bud, (D) Calyx, (E) Staminal corona, (F) Carpels, (G) Stigma top view, (H) Anther with inner corona, and (I) Pollinia. (After Ramakrishna, T.M. et al., The Asclepiadaceae and Periplocaceae of Karnataka, Prasaranga, University of Mysore, Mysore, India, 1995. With permission.)

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Brachystelma maculatum Hook.f., Fl. Brit. India 4: 65. 1883; Gamble, Fl. Madras 2: 851. 1923; Sharma et  al., Fl. Karnataka 164. 1984; S.R.  Srinivasan in Henry et al., Fl. Tamil Nadu Ind., Ser I: Analysis 2: 80. 1987; T.M. Ramakrishna et al., Asclepiadaceae Karnataka 46. 1995; A.P. Jagtap & N.P. Singh, Fsc. Fl. India 24: 186. 1999; Albers & Meve, Ill. Handb. Succ. Pl. Asclepiadaceae 34. 2002. Type: “Canara or Mysore,” Lawson. Etymology: Brachystelma maculatum = spotted Brachystelma; maculatum  = spotted; in allusion to the corolla tube having spots. Stems slender, terete, internodes 2.3–6.7 cm long, glabrous. Latex watery. Leaves opposite, decussate, 6.2–10.5 × 0.1–0.3 cm, linear, long slender almost filiform, apex pointed, base acute, glabrous, lateral veins obscure, midvein prominent; petiole terete, 3–5 mm long, glabrous. Inflorescence axillary, 3–5-flowered umbel. Pedicel filiform, 1–1.5 cm long, glabrous. Calyx 5-lobed, divided up to base, lobes c. 1 × 0.5 mm, subulate, apex acute, glabrous. Corolla tube c. 1.5 mm long, keeled above the middle, lobes five, c. 3 × 0.6 mm, linear, apex subacute or acute, lobes with stiff, purplish hairs. Corona staminal, uniseriate, 5-lobed, c. 1.5  ×  0.2  mm, subulate, opposite the anther lobes much higher than anther, dark brown, glabrous. Stamens 5, c. 1 mm long, pollinia five, pollen masses solitary in each anther cell, yellow, waxy, c. 302.4 × 172.8 µm, with pellucid layer attached by light-brown caudicles. Gynostegium c. 1.5 mm long. Rare in grasslands. Fl.: June–July. Tamil Nadu: Perumal side, Pulney hills (Palni hills), 19-06-1899, A.G. Bourne & E.G. Bourne 2752 (K000820009). India: Karnataka and Tamil Nadu. Endemic. IUCN Category: Rare. Brachystelma mahajanii Kambale & S.R. Yadav, Kew Bull. 69: 9493-2. 2014. Type: India: Tamil Nadu: Nilgiri Distr., Ebbanad village, Kambale & Yadav SSK 49 (Holotype CAL; isotypes SUK! and MH). Etymology: The specific epithet honors a botanist and a teacher, Mr. S.D. Mahajan, Gokhale College, Kolhapur, for his contribution to the conservation of plant diversity of the Northern Western Ghats. Perennial, erect, tuberous herb. Tuber subglobose-discoid, 10–40 mm in diam. Stem is usually one, rarely branched above the ground, erect, 4–9 cm, internodes 0.5–3 cm long, 1–2 mm in diam., puberulous, hairs minute, simple, translucent. Leaves linear-lanceolate, 18–42 × 0.8–1.6  mm, sessile, drooping, fleshy, opposite, decussate, margins ciliolate, apex acute, glabrous except on midrib beneath. Inflorescence extra-­ axillary. Flowers 1–8  at each node, flowers 1–2 in axils of each bract, buds facing upwards, flowers drooping. Bract solitary, 1 × 2 mm, persistent, subulate, hairy, linear. Pedicel 3–5 mm long, terete, pubescent. Calyx 5-lobed, lobes subulate, 1.2 (1.6) × 0.4–0.6 mm at base, acute, puberulous on abaxial surface, glabrous otherwise. Corolla stellate, drooping, c. 3.7–7  cm long, c. 7.5–8 (14) mm in diam., glabrous outside, corolla tube saucer shaped, c. 3 mm in diam., 1–1.2 (1.4) mm deep, abaxially greenish yellow spotted with purple, adaxially yellowish green, glabrous within; corolla lobes 2  (5)–2.5 (5.6) mm long, spreading, rarely reflexed, revolute along margins, glabrous, yellow at upper half, green at lower half, lanceolate, acute at apex, broad at base. Corona biseriate, staminal, and intrastaminal parts fused to a cup-shaped structure; intrastaminal corona 2 × 2  mm, of 5 lobes, lobes undulate, much taller than

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gynostegium, greenish yellow spotted purple outside and within, glabrous; staminal corona lobes linear, triangular at tip, procumbent on stigma. Gynostegium subsessile, anther sacs yellow, stylar head white, flat. Pollinarium: Pollinia yellow with pellucid margin, 0.2 × 0.2 mm, reniform, attached to corpusculum by short caudicle; corpusculum brown. Follicles paired, fusiform, unequal, 5–7.5 cm long, yellowish green, faint purple at maturity; fruiting pedicels straight, c. 1.5 cm long. Seeds 7–9 × 1–1.5 mm, comose, narrowly elliptic, blackish brown at center, margin winged, yellow; coma c. 1.5 cm long, silky white (Figures 2.13 and 2.14).

FIGURE 2.13  Brachystelma mahajanii Kambale & S.R. Yadav: (A) Habit, (B) Inflorescence, (C) Flower: lateral view, (D) Flower: top view, (E) Corona top view, and (F) Seed. (Reproduced from Kambale, S.S. et  al., Kew Bull., 69, 9493–4, 2014. With permission from Trustees of the Royal Botanic Gardens, Kew.)

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FIGURE 2.14  Brachystelma mahajanii Kambale & S.R. Yadav: (A) Habit (November 2012), (B) Habit (March 2013), (C) Close up: inflorescence, (D) Habit (May 2013) & inset: top view of flower, (E) Habit (July 2013), (F–H) Corona: lateral view, and (J) Corona: top view. (Reproduced from Kambale, S.S. et  al., Kew Bull., 69, 9493–4, 2014. With permission from Trustees of the Royal Botanic Gardens, Kew.)

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Eastward slopes of the Nilgiris in Tamil Nadu in soils accumulated on rocks at an altitude of c. 1792 m. Fl.: March–May; Fr.: April–June Tamil Nadu: Nilgiri Distr., Tuber collected from Ebbanad village and grown in SUK Botanic garden, Kambale 204 (SUK). India: Tamil Nadu. Endemic. IUCN Category: Rare. Brachystelma mahendragiriense Prasad et al., Rheedea 27: 135. 2017. Type: INDIA, Odisha: Gajapati district, Mahendragiri hills, 18°58′05.3ʺ N, 084°22′08.6ʺ E, 1393  m, 17.9.2014, K. Prasad & Alok Chorghe 4221a (Holotype, CAL), 4221b–d (Isotype, BSID). Etymology: The new species is named after Mahendragiri hills, its type locality. Herb, c. 20 cm high, perennial. Tuber subglobose, c. 3 cm diam., pale brown. Stems erect, unbranched, c. 1.5 mm thick, pubescent when young, glabrescent on maturity; internodes c. 4 cm long. Leaves opposite-decussate, sessile, linear or linear-lanceolate, 3–6.5 × 0.15–0.2 cm, margins bent inward, apex acute; margins and midrib puberulent. Inflorescence cyme, reduced to two flowers; bract 1, ovate-lanceolate, 1–1.1 mm long, apex acute, glabrous, greenish pink, caducous; bracteoles 2, linear, c. 0.5  mm long, apex obtuse, glabrous. Flowers upright, on 4–6 mm long pedicels, patent. Calyx 5-lobed, sepals 5, triangular-acuminate, 0.9–1.25 mm long, sparsely ciliate, apex acute or obtuse visible between corolla lobes, greenish pink. Corolla rotate, glabrous, yellow, mottled dull/bright pink at base; tube shallow, 0.5–0.75 mm long, mottled pink; lobes rotate, oblongovate, 3.5–4  × 1.4–1.7  mm, margins slightly recurved, apex mucronate with a translucent mucro. Corona biseriate, fused, cupular, 0.5–0.75 × 1.5–1.75  mm, exceeding gynostegium in length, blackish yellow, 5-lobed; upper margins undulate or obscurely 5-lobed, adaxially with horizontally spreading white trichomes in 10 clusters altogether; staminal corona lobes reduced to scales; anther appendages procumbent on stigma head, oblong, c. 4 mm long, apex obtuse, yellowish. Pollinia ovoid, 0.2–0.25 × 0.1–1.5 mm, yellow with pellucid margin along inner side, attached to brown corpusculum by short caudicle. Follicles in pairs or solitary, tapering towards apex, 6–8 cm long; seeds comose, narrowly elliptic, c. 4 × 2 mm, light brown at center. Grass dominated rocky areas, 1350–1400 m. Fl. & Fr.: May–September. India: Odisha. Endemic. IUCN Category: Since not much is known about its population and the species is described based on single location collections, this species is placed under the “Data Deficient” (DD) category (IUCN, 2014). Affinities: Brachystelma mahendragiriense is similar to B. mahajanii, but differs in plants attaining 20 cm height (vs. reach up to 10 cm in B. ­mahajanii); tubers subglobose (vs. tubers discoid in B. mahajanii) leaves never reflexed, margins bent inward (vs. leaves reflexed, margins flat in B. mahajanii) inflorescences 2-flowered at every node, upright (vs. 1–8-­flowered, drooping in B. mahajanii); bract glabrous, greenish pink, caducous; (vs. bract green, hairy at tip, persistent in B. mahajanii); sepals triangular-acuminate (sepals subulate in B. mahajanii); corolla lobes oblong-ovate,

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yellow, mottled dull/bright pink at base, uniformly thick, apex ending into a translucent appendage (vs. lobes ovate, yellow but green at base, broad gibbous base ­distinct from linear thicker upper half, apex acute and devoid of translucent appendage in B. mahajanii); coronal cup is blackish yellow, margins wavy with obscure lobation and with spreading white trichomes at interstaminal positions (corona greenish yellow, blotched purple inside and outside, margins distinctly 5-lobed, glabrous in B. mahajanii). Brachystlma malwanense S.R. Yadav & N.P. Singh, Kew Bull. 48: 59. 1993; A.P. Jagtap & N.P. Singh, Fasc. Fl. India 24: 186. 1999; A.P. Jagtap & S.K. Das Das, Fl. Maharashtra Dicot. 2: 340. 2001. Type: India: Maharashtra, Chowk, about 5 km E of Malwan, S.R. Yadav 4678A (Holotype CAL), 4678B (BSI), 4678 C (K). Etymology: The specific epithet is derived from the type locality Malwan in Maharashtra. Erect, perennial herb, 9–30 cm high, roots subglobose, tuberous. Tuber 2–5 cm in diameter. Stem terete, 0.2–0.4 cm in diam., usually unbranched, hispid. Leaves sessile or subsessile, lanceolate or narrowly elliptic, 4–9  × 0.5–2.5 cm, acute. Flowering axis with internodes 1–3 cm long, bearing flowers in whorls at each node. Flowering nodes with two opposite, subulate, reduced leaves, flowers 2 on either side of each reduced leaf, nodes 3–5 but usually 4-flowered; flowering nodes 3–7. Flowers small, brownish black, bracteate. Pedicels 0.3–0.5 cm long, ­puberulous; bracts subulate, 1–2 mm long. Calyx 5-lobed, lobes 1–2  mm long,  puberulous. Corolla divided almost to base, dark purple and variously variegated within, greenish purple outside; corolla tube short, c. 1.5 mm long; corolla-lobes broad at base, tapering at apex, connate or free, hairy on inner side, 0.6–1 cm long, corolla hairs purple, corolla margins recurved. Corona biseriate, staminal, outer corona c. 2.2 mm in diam., cupular, 5-lobed, lobes hairy, inner corona of 5 dark purple, procumbent processes bent up on stigma, glabrous, entire. Pollinia erect, minute, yellow, attached to brown carriers by short caudicles. Pistil c. 2 mm long, styles indistinct, stigma pentangular. Follicles in pairs, 10–15 cm long, tapering at apex. Seeds comose (Figure 2.15). B. malwanense is an extremely rare, hysteranthous species. Flowering is during February–March while leaves are produced in the rainy season. The species grows in crevices in laterite in the Konkan area around Malwan (Maharashtra) under bushes of Holarrhena antidysenterica (Roth.) A. DC., Carissa carandas L., Ochna obtusata DC., and Lantana camara L. Fl.: February–March; Fr.: April Chromosome number: 2n = 22 (Gosavi et al., 2012) Maharashtra: Pune Distr., Katraj, S.S. Kambale 278 (SUK); Sindhudurg Distr., Kochare, M.Y.Kambale 1128 (SUK). India: Maharashtra, Goa. Endemic. Note: Brachystelma malwanense is similar to B. shrirangii but differs in its generally larger size and its longer flowering internodes, usually 4-flowered nodes, broader leaves, and corolla lobes with purple hairs. B. shrirangii can be readily separated by its much smaller size (under 12 cm high) and other characters given above. IUCN Category: Data Deficient.

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FIGURE 2.15  Brachystelma malwanense S.R. Yadav & N.P. Singh: (A) Plant in vegetative state, (B) Plant in flowering state, (C) Opened flower, (D) Corona, and (E) Pollinarium. (Reproduced from Yadav, S.R. et al., Kew Bull., 48, 59–61, 1993. With permission from Trustees of the Royal Botanic Gardens, Kew.)

Brachystelma matthewianum P.V. Bruyns & S.J. Britto, Haseltonia 22: 51. 2016. Type: India, Tamil Nadu, west of Madurai, 1150 m, Bruyns 11488 (Holotype E). Etymology: This species was named in honor of Dr. Koyapillil M. Matthew (1930– 2004), who for many years was the director of the Rapinat Herbarium at St Joseph’s College, Tiruchirapalli, and who initiated and published the well-known illustrated Flora of the Tamil Nadu Carnatic.

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Perennial geophytic herb to 5 cm tall, arising from an underground globose to a flattened pale brown tuber, 1–2 × 1–2.5 cm. Stem annual, few-branched, spreading, slender, dark greenish purple, terete, sparsely puberulous, internodes 0.5–1 cm long. Leaves nearly sessile, slightly fleshy, opposite, linear, sparsely puberulous, 1–3 × 0.1–0.5 cm (those at base reduced to subulate scales to 2 mm long), with cuneate or subacute base, margins entire and shortly ciliolate, gradually acute, apiculate. Inflorescence in few extra-axillary cymes alongside middle to upper nodes, 1–3-flowered, flowers opening in gradual succession; sessile, sparsely puberulous; pedicel 1–5 mm long, slender, puberulous; bracts linear, c. 1 mm long; sepals lanceolate, green, 1–1.5 × 0.5 mm, sparsely puberulous. Corolla campanulate, outside purple-brown, glabrous; inside pale yellow becoming brown near tips of lobes, glabrous; tube c. 1.5 mm long and 2 mm broad at mouth, cup-like and equaling the corona; lobes erect to spreading, deltate with slightly reflexed margins, 3.0–3.5 × 2 mm, acute. Corona 2-seriate; outer series forming cup-like structure 1.5 mm tall and 2 mm broad, filling tube and much exceeding the height of the anthers, consisting of five erect and truncate lobes heavily indented between them with erect rims, slightly less than 1 mm wide, glabrous outside and with straight white hairs inside in patches alternating with anthers, mottled and margined with purple-black on yellow; inner series arising from below the middle of the outer series, of 5 small deltate lobules just touching backs of anthers, c. 0.25 mm long, yellow mottled with purple, glabrous (Figures 2.16 and 2.17). Brachystelma matthewianum is so far known only from a single area on the eastern slopes of the higher mountains west of Madurai, which form outliers of the Western Ghats. Here, it occurs in small, almost level patches of shallow soil overlaying granite on steep slopes. These patches have relatively little other vegetation except for other small geophytes and some small annuals, while deeper soil nearby and higher up the slopes has an often-dense covering of trees. Fl. & Fr.: August– India: Tamil Nadu. Endemic. IUCN Red list Category: Brachystelma matthewianum is known only from the type locality, where few plants were seen: rated Vulnerable (V,  IUCN, 2010). Note: Brachystelma matthewianum is also a non-climbing plant. It is particularly small, rarely exceeding 5 cm tall, with short stems that branch sparsely and remain close to the ground. Most of the flowers are borne before the leaves are fully developed. Along with B. rapinatianum, it  shares the non-climbing habit and significant corolla tube, but this is shorter than in B. rapinatianum and only half the length of the lobes. In  B. matthewianum, the deeply cup-like corona formed by the considerable fusion of the outer series of lobes is also suggestive of B. mahajanii. However, in B. matthewianum, the cup-like structure formed by the outer lobes is taller and has strongly undulating sides (deeply indented behind the anthers). Unlike B. mahajanii, where they are lanceolate, the corolla-lobes here are deltate, while the corona is just contained in the corolla tube (whereas it projects from a short tube 1–1.2 mm long in B. mahajanii). Inside, the cup-like outer corona has patches of white hairs alternating with the anthers and these are absent in B. ­m ahajanii (Bruyns and Britto, 2016).

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FIGURE 2.16  Brachystelma matthewianum Britto & Bruyns: (a) Habit with tuber, (b) Flower front view, (c) Corona, (d) Pollinarium, and (e) Follicle. (Courtesy of Karuppusamy.)

Brachystelma megamalayana Karupp. sp. nov. Type: India, Tamil Nadu, Theni Distr., Megamalai Wildlife Sanctuary, 20-07-2014 S. Karuppusamy & V. Ravichandran 1025 (holotype MH; Isotypes SGH). Perennial, erect, tuberous herb. Rootstock globose to subglobose, tuber 30–60 mm diam. Stem usually two from each tuber, erect, 5–10 cm tall, unbranched, fleshy, 1.2 mm diam., 4–5 internodes, pinkish, finely puberulous, hairs translucent. Leaves simple, opposite, linear-lanceolate, 30–60 × 5–20 mm, variable in size, acute apex, margin recurved, minutely ciliolate along the margin and midrib, pinkish, pubescent beneath, sparingly tomentose above; mid-rib puberulous beneath, narrowly grooved above; petiole absent or very short, 0.1 mm long; first pair of leaves fleshy,

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FIGURE 2.17  Brachystelma matthewianum Britto & Bruyns: (a) Habit with tuber, (b) Flower-front view, and (c) Tuber. (Courtesy of Karuppusamy.)

linear-lanceolate, pinkish, pubescent, short,  up to 1.5 cm long. Flowers in sessile umbels, axillary or crowded on nodes, 2–6-flowered, drooping; pedicel stout, cylindrical, 0.8–1 cm long, pinkish, puberulate tomentose. Bracts linear, solitary, pinkish, 10–13 mm long, puberuous, persistent; bracteoles 3 mm long, linear, purplish, puberulous. Calyx deeply 5-lobed, lobes 0.5 × 0.2 mm, ovate-lanceolate, fleshy, pinkish, puberulous outside, glabrous inside. Corolla lobes 5, broadly ovate at base, linear-lanceolate above half, acute at apex, tips incurved, 1 × 0.5 cm, rarely spreading, puberulous tomentose outside, glabrous inside, varigate by yellow, brown, greenish yellow to pinkish brown, rarely dark purple blotches inside, sometimes dark purple or brownish above half, mid-vein prominent outside; corolla tube campanulate, 0.4 mm long, 5 mm wide, pale brown to pinkish blotches outside. Corona biseriate, outer corona dark purplish to brownish, cupular, 5-lobed; lobes raise above corolla tube, orbiculate to ovoid, 0.7 × 0.3 mm; lobes minutely truncate at the apex, undulate; tufts of dark purple ciliate hairs on inner face of the lobes; inner corona 5-lobed, lobes subulate, linear, glabrous, procumbent on the anther column, yellowish in color,

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slightly truncate at the apex. Gynostegium sessile, anther sacs yellow, stylar head white, flat. Pollinarium: Pollinia 5, erect, 0.3 × 0.2 mm, yellow with pellucid margin, oblong, attached to the corpusculum by 0.2 mm long caudicle; ­corpusculum brown to pinkish, triangular. Mericarps paired, 0.3 mm long, double, subequal, erect, yellowish green, pinkish at maturity, 5–7 cm long, linear-cylindric,  glabrous, divergent, apex blunt, 7–12-seeded; fruiting pedicel erect, 1 cm long, stout, c­ ylindrical, pinkish,  puberulous. Seeds elliptic-ovate, brown, 10 × 0.2 mm, with central dark brown embryonic region, margin winged, light brown; coma silky, 2.5  cm long (Figures 2.18 and 2.19). Fl. & Fr.: June–August. Rcongnition: Brachystelma megamalayana resembles B. mahajanii (Kambale et al., 2014) in habit, by its other characters approaches to B. matthewianum (Britto

FIGURE 2.18  Brachystelma megamalayana Karupp.: (a) Habit with tuber, (b) Flower lateral view, (c–e) Flowers front view, (f) Corona, (g) Pollinarium, (h) Follicle, and (i) Seeds with coma. (Courtesy of Karuppusamy.)

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FIGURE 2.19  Brachystelma megamalayana Karupp.: (a) Habit, (b) Plant with tuber, (c) Flowers with inflorescence, and (d) Dehiscing follicle. (Courtesy of Karuppusamy.)

and Bryun, 2016); from the former species it differs in double shoots from each tuber, short stem, 2–6-flowers at each node, variously colored corolla lobes, tufts of hairs on outer corona lobes with truncated top and differs from the B. matthewianum in the tall habit with larger tuber, long semisucculent leaves, 2–6-flowers in cymes, longer pedicel, yellow-colored corolla lobes, wider corolla tube, and shape of the corona. Distribution: India: Tamil Nadu; Theni District, Megamalai Wildlife Sanctuary. Conservation status: Brachystelma megamalayana so far known from the type locality only. About 224 individuals were located. According to IUCN criteria, the species falls under the category Data Deficient (DD) (IUCN, 2016.2). Brachystelma nallamalayana Prasad & Rao, J. Threatened Taxa 5: 4904. 2013; Sadasivaiah et al., Asian J. Plant Sci. Res. 6: 27. 2016. Type: India: Andhra Pradsh, Bheemudikolanu, Nallamalai in Eastern Ghats, BR & KP 34667 A (Holotype), 34667 B-D (Isotype BSID). Etymology: The new species is named after the type locality, the Nallamalais of Andhra Pradesh, India.

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Erect herbs, up to 80 cm high. Tubers in varied shapes, generally fusiform, to 6 × 3 cm diam., brownish. Stem solitary, terete, 1–2 mm in diam., unbranched, ridged, succulent, glabrous, glaucous green; internodes up to 10 cm long. Leaves opposite, decussate, fleshy, sessile; linear, up to 19 cm long and c. 2 mm diam., margin undulate, apex acute, glabrous, sometimes hairy on the mid vein. Inflorescence crowded at apical nodes, lateral, shortly peduncled, umbellate, 5–6-flowered, pendulous. Pedicels terete, filiform, up to 1.3 cm long, glabrous. Bracts and bracteoles persistent, linear, acute at apex, glabrous, greenish pink; bracts 2 × 0.5 mm; bracteoles 1 × 0.2 mm. Calyx 5-lobed, united at base; lobes linear, 3 × 0.5 mm, acute at apex, 3–5 veined, glabrous, pinkish green; corolla 5-lobed. Corolla tube up to 1mm long, shallow; corolla lobes erect, linear, 5–9 × 2  mm diam., acute but appears obtuse due to inturned apex, faintly 5-veined, margins involute with 1–2 mm long pink hairs throughout, more dense at apex; lobes basally pale white with black spots and yellow above without any blotches. Corona c. 3 mm across, biseriate, glabrous; interstaminal corona obscurely 5-angled, cupular, forms a continuous ring around the gynostegium, brown; staminal corona 5-lobed, black; lobes appressed to the back of the anthers, exceeding and hooded on the style apex. Pollinia waxy, yellow, globose, c. 200 μm long, margins pellucid at apex, attached by light brown tubular caudicles to a redcolored corpuscle. Style apex pentangular. Follicles paired, linear, c. 6 × 0.3 cm, glabrous, glacous. Seeds  comose, 5–7 × 3–5 mm, ovate-oblong, 15–20 in each follicle, light brown at center (Figures 2.20 and 2.21). This species occurs amidst grasses on open hillslopes of dry deciduous forests at an altitude between 550 and 600 m, associated with Cymbopogon coloratus and Heteropogon contortus in the Nallamalai Hill ranges. Fl.: July–August Talangana: Way to Saleswaram (MBNR), B. Sadasivaiah & B. Kalpana, 2073; Way to Srisailam, Near Rasamola Baavi (MBNR), B. Sadasivaiah & K. Prasad, 2142 (Govt. Degree College, Wanaparthy). India: Andhra Pradesh, Telangana. Endemic. IUCN Category: Brachystelma nallamalayana Prasad & Ravi Prasad Rao has restricted distribution and currently is known only from Nallamalais. Until further explorations determine its distribution range, biology and threats, the species cannot be assessed. While the available information perhaps qualifies it initially as a Critically Endangered species, it is at present Data Deficient (Prasad and Rao, 2013; Sadasivaiah et al., 2016). Brachystelma naorojii P. Tetali et al., Rheedea 8: 75. 1998; A.P. Jagtap & N.P. Singh, Fasc. Fl. India 24: 187. 1999; A.P. Jagtap & S.K. Das Das, Fl. Maharashtra Dicot. 2: 341. 2001; M.R. Almeida, Fl. Maharashtra 3A: 225. 2001. Type: India, Maharashtra: Hillslopes of Gavadewadi near Pandav Dhara, 3  km from Shindewadi Phata on Pune—Satara National Highway (NH-4), P.Tetali 72 A (Holotype—AHMA!); P. Tetali 17895, 17896 (Paratype—K). Etymology: The species has been named after Mr. Naoroji Godrej, a noted industrialist, to commemorate his keen interest in plant conservation. Vern.: Marathi: Haman, Hamman batata A slender perennial herb, 30–55 cm high. Rootstock tuberous, round, or ovate when young, flattened in mature plants, fleshy, 7–9 × 4–6 cm. Stem glaucous

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FIGURE 2.20  Brachystelma nallamalayana Prasad & Rao: (A) Habit, (B) Flower, (C) Bract, (D) Bracteole, (E) Calyx, (F) Corona top view, and (G) Pollinia. (After Prasad, K., and Rao, B.R.P., J. Threatened Taxa, 5, 4904–4906, 2013.)

green, occasionally tinged with purple, usually unbranched with deltoid leaf scales, sparingly pubescent, occasionally with appressed hairs. Latex watery, yellowish green, bitter. Internode between first and second pair of leaves 5–30 cm long, the first pair of leaves always small. Leaves sessile, opposite-decussate, lanceolate, ovate-oblong or ovate-lanceolate, 1.5–9 × 0.5–4 cm, glabrous above and pale beneath, apex acuminate, margin ciliate, hairs swollen at base. Flowering nodes with opposite, 2–3 mm long scale leaves. Flowers bracteate, two on either

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FIGURE 2.21  Brachystelma nallamalayana Prasad & Rao: (A) Habit, (B) Bud, (C) Inflorescence, (D) Flower, (E) Tuber, and (F) Fruit. (After Prasad, K., and Rao, B.R.P., J. Threatened Taxa, 5, 4904, 2013.)

side of the scale leaf, green turning to brick red, reddish violet, and finally to reddish purple—darker and puberulous within, hairs white. Pedicel 5–7 mm long, sparsely hairy. Calyx 5-lobed, lobes 5, 1.5–2 mm long. Corolla deeply 5-fid, lobes 0.9–1  cm long, thickly hairy on the inner side. Corona cupular, 5-lobed, lobes with long white hairs, margin dark purple, lower portion of the corona white, 2.3–3 × 3 mm, sparsely hairy, obtusely five-angled, coronal lobes produced into

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5 inner lobes incumbent and adpressed on the stamen. Stamens 5, pollinia globose, yellow, 405 µm long, style-apex pentagonal. Follicles  single or in pairs, 6 cm long, tapering at the apex. Seeds comose, dark brown with light brown margin, 0.8–1 cm long, coma silky white, up to 2 cm long. Rare on partially degraded hillslopes and open hilltops among grasses in Maharashtra. Fl. & Fr.: May–June Maharashtra: Sindhudurg Distr., Shirwal, D.K.Kulkarni s.n. (AHMA); Pune Distr., Shindewadi, M.Y. Kamble 1172 (SUK). India: Maharashtra. Endemic. Note: This species is said to be allied to B. kolarensis Arekal & Ramakrishna from which it differes in having larger tubers and leaves, flowers without purple spots, large cupular corona, with dark purple lobes and bigger pollinia. Its other distinguishing feature is that it flowers in summer. IUCN Category: Data Deficient. Brachystelma nigidianum Raja Kullayisw. et  al., Kew Bull. 71: 1. 2016 & Fl. Thummalapalli Uranium Mining Area 533. 2017. Brachystelma volubile auct. non Hook. f.; Kullayiswamy et al., Rheedea 22: 107. 2012. Type: India: Anantapur Distr. Nigidi forest, K.Rajakullayiswamy 41853 (Holotype SKU; Isotypes BSID, CAL, K, MH). Etymology: The species was named after the type locality Nigidi forest, which is a part of the Eastern Ghats, Anantapur district of Andhra Pradesh state, India. Perennial climbing herb, to 1.5 m long. Tuber discoid-subglobose, 1 or 2 subterranean, brown, c. 2.8 × 3.5 cm; stem herbaceous, terete, glabrous, sometimes branched. Leaves linear-lanceolate, 5–16 × 0.2–0.5 cm, narrowed into 3–5 mm long petiole, base acute, margin entire, apex acuminate, scabrous above, hirsute beneath on raised midrib, m ­ argin ciliate, caducous, leaves reduced towards apex on stem, sessile c. 3 mm long, scabrous. Petiole scabrous, canaliculate with brown colleter (gland) at base. Inflorescence extra-axillary, 2- or 3-flowered; peduncles 5–6 mm long, glabrous. Bracts lanceolate, glabrous, c. 4 mm long; ­bracteoles lanceolate, glabrous c. 3 mm long. Pedicels c. 9 mm long, glabrous, purplish green. Calyx 5-lobed, lobes linear-lanceolate, subulate, 9–10 mm long, glabrous, purplish green. Corolla 3.3–3.5 cm long, c. 4.4 cm across (when open); corolla tube c. 9 mm long, urceolate or urn-shaped, base, middle, and mouth of corolla tube c. 9 mm, c. 7 mm, and c. 11 mm wide, respectively; glabrous, greenish with brown spots without, and slightly ribbed, pubescent with fine hairs, dark brown with light yellow lines within; lobes deltoid at base (two times wider than long) tapering toward apex, c. 2.6 × 0.3 mm, cohering at apex when young and free when mature, acuminate at apex, glabrous, greenish outside, reddish maroon, densely pubescent within, scattered pilose along margin (hairs c. 2 mm long), margin slightly outcurved, yellow striations at base of lobes. Corona c. 2 mm high, c. 4 mm across; outer corona cupular, reddish maroon, c. 2 mm across, pubescent, 5-lobed, each lobe slightly bifid; inner corona 5-lobed, dorsiventrally flattened, c. 1 mm long, basally fused with outer ring, lobes 5, free, oblong-ovate, yellow with maroon tinge at middle, apex obtuse, overlapping anthers with pollen chambers below. Pollinaria c. 0.5 × 0.5 mm across; pollinia ovoid, c. 0.4 × 0.3 mm, yellow, basally attached by hallow hyaline caudicles to a brown corpuscle, pollinia

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with germination crests towards apex more than half as long as ­pollinium; ovary c. 2 mm long, style short. Follicles in pairs, slightly fusiform, 5.5–7.5 cm × 2.5–4 mm, tapering, connate at apex, slender, smooth, glabrous, pale green. Seeds flattened, ovate, c. 5 × 2.5 mm, light brown, outer margins thin, comose; coma 9–14 mm, white silky (Figures 2.22 and 2.23). Rare in Nigidi forest in Anantapur district of Andhra Pradesh. Fl. & Fr.: October–December India: Andhra Pradesh. Endemic. IUCN Category: Brachystelma nigidianum is known from the type locality with only 10 individuals. The species is therefore assessed as Data Deficient (DD) according to IUCN version 3.1 (2001). Brachystelma parviflorum (Wight) Hook.f., Fl. Brit. India 4: 65. 1883; Albers & Meve, Ill. Handb. Succ. Pl. Asclepiadaceae 38. 2002. Eriopetalum parviflorum Wight Contrib. Bot. India 35. 1834. Type: A single specimen is available at K (K000820014), which was collected from Madras, Tamil Nadu by Wight. Etymology: Brachystelma parviflorum: small-flowered Brachystelma; parviflorum = small- (or little-) flowered; in allusion to the flowers being small (