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Volume I of Southern Africa and Madagascar Peter V. Bruyns 2005 Stapeliads UMDAUS PRESS P.O. BOX 11059 HATFIELD 0028 SOUTH AFRICA E-Mail: [email protected] Website: www.sukkulents.net FIRST PUBLISHED 2005 © Peter V. Bruyns, Bolus Herbarium, University of Cape Town, South Africa © Photographs: Author and others mentioned in the acknowledgements. © Illustrations: Peter V. Bruyns PRODUCTION Alex Fick and Kotie Relief DESIGN Tersia van Rensen COLOUR SEPARATIONS Jason Pyper PRINTING Tien Wah Press, Singapore BINDING OF SPECIAL EDITIONS Peter Carstens All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner(s). Sponsors Edition: Vol. I - ISBN 1-919766-33-2. Vol. II - ISBN 1-919766-34-0 Collector’s Edition: Vol. I - ISBN 1-919766-35-9. Vol. II - ISBN 1-919766-36-7 Standard Edition: Vol. I - ISBN 1-919766-37-5. Vol. II - ISBN 1-919766-38-3 II VOLUME I: Acknowledgements ................................................................................................................................ IV Dedication ………………………………………………………………………………………….. V New combinations published in this work ………………………………………………………… VI Introduction ………………………………………………………………………………………... 1 Historical Sketch …………………………………………………………………………………... 2 Classification of the Stapeliads ……………………………………………………………………. 4 Relationships among the genera ……………………………………………………………….. 5 Species concepts among the Stapeliads………………………………………………………….. 7 Morphology of the Stapeliads ........................................................................................................... 9 The plant.......................................................................................................................................... 9 The flower.................................................................................................................................... 16 Fruit, seed and seedlings ............................................................................................................. 31 Chromosome numbers ................................................................................................................ 33 Pollination Biology of the Stapeliads.............................................................................................. 34 Flowering times ........................................................................................................................... 34 The mechanics of pollination in the Stapeliads and self-fertility................................................. 34 Pollinators and attractants……………………………………………………………………….. 35 Biogeography of the Stapeliads....................................................................................................... 39 Patterns of distribution and diversity……………………………………………………………. 39 Habitat and ecology....................................................................................................................... 46 Cultivation of the Stapeliads............................................................................................................ 53 Propagation…………………………………………………………………………………….. 53 Diseases........................................................................................................................................ 55 Uses of the Stapeliads...................................................................................................................... 58 Systematic account ......................................................................................................................... 59 Key of the genera in Southern Africa and Madagascar………………………………………… 59 1. Australluma…………………………………………………………………………………….. 61 2. Bayncsia ..................................................................................................................................... 66 3. Duvalia…………………………………………………………………………………………. 92 4. Hoodia ........................................................................................................................................ 92 5. Huernia........................................................................................................................................ 130 6. Larryleachia................................................................................................................................. 212 7. Lavrina………………………………………………………………………………………….. 229 8. Notechidnopsis………………………………………………………………………………….. 232 9. Ophionella.................................................................................................................................... 234 10. Orbea.......................................................................................................................................... 240 VOLUME II: 11. Pectinaria.................................................................................................................................... 331 12. Piaranthus................................................................................................................................... 345 13. Quaqua....................................................................................................................................... 369 14. Richtersveldia ............................................................................................................................ 416 15. Stapelia....................................................................................................................................... 418 16. Stapelianthus .............................................................................................................................. 490 17. Stapeliopsis................................................................................................................................. 507 18. Tavarcsia .................................................................................................................................... 526 19. Tritentea………………………………………………………………………………………... 531 20. Tromotriche................................................................................................................................. 551 21. Hybrids ....................................................................................................................................... 575 Appendix .......................................................................................................................................... 579 Insufficiently known names ............................................................................................................. 579 Possible hybrids ............................................................................................................................... 580 References ........................................................................................................................................ 582 Index to scientific Stapeliad names................................................................................................... 589 List of Subscribers............................................................................................................................. 600 III Contents This work would not have been possible without the help generously given to me over many years by many individuals. Encouragement in the pursuit of this fairly esoteric interest came from many sources, but it was particularly fostered by excursions into the field with Walter Wisura, formerly the curator for succulents at Kirstenbosch, and by M. Bruce Bayer, formerly the curator of the Karoo Botanic Garden, Worcester. Kind and helpful farmers have allowed access to their properties to look for stapeliads. The generosity that I have experienced on farms in South Africa and Namibia is unforgettable. I particularly wish to thank the following for their hospitality and help: Auriol Batten, John and Susanne Bell, Jossie and the late Ortwin Brandt, Johann Bouwer, Dennis and Antjie de Kock, the late Gesina Carolina (Tant Gesie') du Plessis, Johann and Desiree du Toit, Charlotte Grabow, David and Dorothy Green, Gwynne Griffiths and Chris Milne, Norbert Hahn, Adolf Klein, Petrus and Elizabeth Kotze, Hermann Kunert, Erica and Leo Latti, Molly, Chris and Marina Lochner, Daan and Pat Marais, Chris and Elna Marincowitz, Bernt and Gerrie Maritz, Ralph and Christiana Peckover, Shirley Pienaar Nico and Cheryl Pretorius the late Jaap and Kila Snyman, Ken and Lorraine Tarr, Steven Theron, Lucy van der Vyver, Pieter van der Westhuizen, Gebhard von Alvensleben, Buys and Margaretha Wiese, Gordon and Ada Whittal and Mirinda Wilken. The loan of slides by G.D. Court, D. de Kock, H.C. Kennedy, W.R. Liltved, J.A. Retief and G.D. Tribe is gratefully acknowledged, as is the logistical support provided by my brother John in Botswana. D. de Kock, E. Heunis and H.C. Kennedy have provided generous assistance by growing plants for me and allowing these to be removed for photographs and the making of herbarium specimens. I have also grateful to Colin Walker for copying literature for me on various occasions and especially to David Goyder for his willingness to answer frequent enquiries and to search for many obscure references. Christiane Anderson (Michigan University) also helped significantly towards sorting out the complex synonymy of Orbea variegata and in locating some obscure publications. David Richards and Margaret Sandwith helped with several biographical facts about collectors and Callan Cohen with some useful references. I also wish to express my gratitude to Trevor Sewell and the staff of the Electron Microscope Unit at the University of Cape Town for their ever-willing assistance. The University of Cape Town has also provided funding for my research. Missouri Botanic Garden granted permission for the reproduction of figures already published. I would like to thank Umdaus Press for undertaking the publication of this work. Finally, I wish to thanks my parents for their support and encouragement over many years and Cornelia for her encouragement and for her useful suggestions, which have helped to improve this work substantially. IV Acknowledgements V For my family, Bruce and Steven Australluma ubomboensis Duvalia caespitosa subsp. pubescens D. caespitosa subsp. vestita Huernia barbata subsp. ingeae H. blyderiverensis H. guttata subsp, reticuiata H. hislopii subsp. cashelensis H. hystrix subsp. parvula H. verekeri subsp. angolensis H. verekeri subsp. pauciflora H. zebriana subsp. insigniflora Larryleachia cactiformis var. feline Orbea longii Pectinaria longipes subsp. villetii Piaranthus cornutus var. ruschii P. geminates subsp. decorus P. geminates subsp. framsii Stapelia grandiflora var. conformis S. hirsute var. baylissii S. hirsuta var. gariepensis S. hirsuta var. tsomoensis S. hirsute var. vetula S. paniculata subsp. kougabergensis S. paniculata subsp. scitula Tromotriche pedunculata subsp. longipes VI New combinations published in this work he stapeliads are the most highly succulent members in the tribe Ceropegieae and belong to the subfamily Asclepiadoideae of the family Apocynaceae. They are all fleshy stem-succulents that are more or less totally without leaves and only rarely have thorns. They exhibit an extraordinarily wide range of flower shapes and sizes (with some of the largest I lowers in the plant kingdom found among them) and there is also a wide range of complicated strictures in the centre of the flower hat are associated with the process of pollination. The flowers are specialized exclusively for fly pollination and this diversity appears to have arisen in response to the wide spectrum of sizes of flies that are present in the region combined with the wide range of geological and topographical niches in the area. These volumes aim to document the extent of this diversity for southern Africa. The last detailed treatment of the stapeliads wTash e Stapelieae by Alain C. White & Boyd L. Sloan, ewhich appeared in three volumes in 1937. This monograph, with a total of over 1200 pages, covered the whole group in unprecedented detail and today copies are valuable and much sought after. However, much exploration has taken place in the 67 years sincTe he Stapelieaea ppeared. This has led to discovery of many new species and the realization that many of the ‘species’ discussed by White & Sloane, who never had the opportunity to see any of these plants in their natural habitat, were not species at all. Therefore, despite its pre-eminence, The Stapelieae is now considerably out-of-date and is more or less impossible to use for the identification of recent collections. The need for a replacement is consequently quite urgent. At present the stapeliads consist of, in total, 326 species. These volumes are the first attempt to present a renewed account of them and they deal with the southern African species. For present purposes, southern Africa is taken to be that portion of Africa which lies south of 17°S. Here we include the whole of Namibia, Southern Africa, Botswana and also all of Zimbabwe and Mozambique. The island of Madagascar is (perhaps rather unconventionally) also included, but this is done since the few stapeliads that grow there are more closely related to others in southern Africa rather than to species from further north in Africa. At present the number of stapeliad species that occur in this area is 182. These 182 species are distributed in 19 genera and all but four of these genera are endemic to the region. Of the 182 species found in southern Africa, all except 15 are endemic to this area and these 15 non-endemic species mostly extend slightly beyond the borders of our area into Angola, Malawi and Zambia. This means that 92% of southern African stapeliads are endemic to the region. Thus these volumes will deal with somewhat more than half of the total number of known stapeliads and will have little overlap with any account of the species from further north. This works brings together, from many disparate sources, the results of the exploration and research that has taken place over the past 60 years on southern African stapeliads. It consists of two sections, the first of which is introductory and gives the leader some conception of the complexity and diversity that has evolved in the group, especially in the floral structures. There are considerable numbers of terms that are peculiar to the study of asclepiads in the general and stapeliads in particular and these are explained and illustrated with examples in the section on morphology. The second section is the systematic account, which contains details of the individual species. Keys are provided to all the genera, to all the species and to all the subspecies and varieties. All 182 species are described and discussed. Each species is illustrated by means of several color photographs, with a map showing its distribution and with line drawings in which some of the essential, but minute details of the flowers are highlighted. In the photographs, particular emphasis is placed on showing the variability of the species. Vouchers for the PVB numbers cited in the text are in the herbaria BOL, NBG, PRE, K and MO. Where latitude and longitude 'grid squares' are cited in the text this follows the system of Edwards & Leistner (1971). 1 T Introduction hile there is no doubt that stapeliads have described in detail by Brown in his Stapeliae been known in southern Africa for as long as Barklyanae( Brown 1890). people have been in the area, they were first During his preparation of the accounts noticed by Western travelers in 1624 when Justus of the Asclepiadaceae for the Flora Capensis Huernius found what is today known aOs rbea variegata (Brown 1907-09), Brown also encouraged on the slopes of Table Mountain. By 1700 Stapelia Charles Eustace Pillans, who was a hirsuta was known, and Quaqua incaenata and Q. distinguished civil servant in the government mammillaris had also been discovered prior to the of the time in South Africa and his son Neville publication of Linnaeus's Species Plantarum in 1753, Stuart Pillans (fig. 1), a botanist employed at although Linnaeus listed only Stapelia variegata and the Bolus Herbarium in Cape Town, to gather S. hirsuta in this account. At this stage exploration of material. This they did with tremendous success and the then relatively unknown interior of southern Brown ended up naming eight species in their honor, Africa was very tentative and fraught with all manner one in nearly every genus of southern African of dangers, both real and imagined, and this stapeliads. They appear to have travelled by rail and considerably hampered botanical discovery in the stopped at some of the small sidings of the Karoo to region. The documentation of stapeliads of southern make extensive collections around them, presumably Africa can really be said to have begun in earnest residing in the area for a few days. In this way they with the explorations of C.P. Thunberg and Francis covered extensive parts of the Great Karoo. In 1926, Masson. Thun-berg spent nearly three years at the N.S. Pillans also made an important pioneering Cape and during this time he and Masson explored journey, part of it by ox-wagon, to the Richtersveld some areas together. Masson spent a total of about and on this occasion several species were discovered, 12 years at the Cape during two visits to the colony among which was the singular and rare Stapeliopsis and left for the last time early in 1795. He was sent neronis. To a large extent they completed the basic to the Cape by the king of England, who acted on the exploration for stapeliads of the Cape Province and it advice of Joseph Banks. Banks had been on the first could be claimed that only sporadic and relatively minor voyage around the world led by Captain Cook and, Fig. 1 . Neville S. Pillans (courtesy University of Cape discoveries have been made since then. Town) when the ship put in briefly at the Cape, had been very It was shortly after N.E. Brown's death in 1934 much impressed by the richness of the flora there. that The Stapelieae appeared (White & Sloane 1937), During his two sojourns at the Cape, Masson between 1791 and 1793, and it was first detected the monumental and now famous publication of Alain on the European mainland (in southern Spain) introduced many novel and remarkable species to Campbell White (1880-1951) and Boyd Lincoln before 1822 by Mariano Lagasca y Seguro Europe, has he send both live plants and seeds back to (Bruyns 1987b). Sloane (1886-1955). White was a wealthy person by London. He seems to have had a particular Stapeliads continued to be discovered inheritance (his father having made his fortune in New predilection for stapeliads and described 41 species in intermittently in southern Africa after the York real estate), educated in languages, who his book Stapeliae Novae, of which most were new in departure of Masson. However, it was the entertained a lasting passion for chess and, later on, also the sense of the time. He cultivated these plants in his enthusiastic encouragement and enquiries for certain succulents. Sloane, who was a teacher and garden at Cape Town and took several of them back of N.E. Brown at Kew that really stimulated principal of various middle schools, mainly in further interest in them at the Cape Colony. to London with him, thereby introducing them into Brown worked as a botanist at Kew from California, was greatly interested in cacti and cultivation in Europe. 1873 until his retirement in 1914 (after which succulents and had a long-time involvement with the It was around this time (in the latter half of the he continued there unofficially until his death American Cactus and Succulent Society, which 18th century) that stapeliads first began to be in 1934) and he was particularly interested included being president of the society in 1931 and discovered in other arid parts of the so-called Old in succulents, especially the succulent 1932 (Mitich 1994). They first met in December World. Pehr Forsskal collected five stapeliads in the Aizoaceae and stapeliads. An unusual trait for 1930 and, after White moved to Pasadena, a botanist was that he cultivated many of them Arabian Peninsula during an ill-fated journey in about California, in 1931, he built up a very large private personally. This enabled him to gain a greater 1762. Species became known from India in about understanding of them and he also observed collection of stapeliads. Soon they decided to 1790 when William Roxburgh first documented and illustrated their flowers in minute detail collaborate on a bock stapeliads and this was Caralluma adscen-ciens, while Henrietta Antonia under the microscope, producing a great published at White's expense in September 1933 Clive collected C. umbellata around the turn of the many very accurate and extremely attractive (White & Sloane 1933). Their first effort was so century. It appeared, therefore, to the botanical drawings. At Brown's instigation, considerable well received that White decided to expand it to a community of Europe that stapeliads were very numbers of stapeliads were sent back to Britain comprehensive monographic treatment and this by Henry Barkly during the period 1870-7, exotic and naturally occurred only in hot, arid, far- appeared in three volumes in February of 1937, again while he was governor at the Cape. With off lands, where much danger and mystery sur- much new exploration coming alongside the produced at his own expense. After this, together with rounded them. Consequently, the discovery of similar discovery of diamonds deep in the interior of the South African botanist R.A. Dyer, they produced peculiar plants on the southern shores of Europe itself South Africa, more stapeliads were collected two volumes on the southern African representatives by Giovanni Gussone in 1828 caused considerable well away from Cape Town than had been the of the Succulent Euphorbieae (White et el. 1941). surprise in certain circles. Gussone collected what case before. Barkly seems to have collected Further volumes as sequels to these two were planned, later came to be known as Caralluma europaea, relatively few of these himself, but he certainly but White had already moved away from California encouraged travelers who noticed them to although the Danish botanist and traveler Peder K.A. in 1937 and never completed them (Sloane 1952). bring him material. He was able to amass an Schoesboe had been the first to notice it in Morocco impressive collection of them and these were Subsequent to the efforts of Pillans, it was 2 W Historical Sketch Fig. 2. Carl A. Lückhoff as a medical student in about 1934. (courtesy Marga Hoffmeyr). particularly the collecting forays into Nama- From 1900 until 1914 Dinter was employed by the In areas outside South Africa and Namibia qualand and the Karoo by Charles Theodore German Colonial Government to document the stapeliads are generally rarer and consequently there de Mornet Villet and his wife, Elizabeth, vegetation of Namibia and investigate certain have seldom been collectors or botanists who have that resulted in the discovery of several problems that arose in the rural areas. During this specialised in their study. In Zimbabwe they new species between 1928 and about 1940. period he often had at his disposal a railway carriage appeared very sporadically among the collections of Other species became known as new roads which would be parked at a siding to enable him to botanists and others. This changed once the amateur gave collectors greater access to previously botanise in the area. He covered much of the less botanist Leslie Charles (Larry) Leach (1909-96) gave impenetrable mountainous areas. Many of accessible territory, such as the Namib Desert south up his business activities to devote all his time to these were brought to Carl August Lückhoff of Lüderitz and the Great Karas Mountains, by foot taxonomic studies of stapeliads, succulent (fig. 2), who was a medical practitioner and, or accompanied by an ox-wagon. It is only in areas Euphorbiaceaea nd Aloe. Leach had been in the army in for a long time, the chief medical officer for that Dinter never visited, for example, the Tiras Britain, the land of his birth, and after encountering the South African Life Insurance Company. He was Mountains, the Kaokoveld and the region east of problems there he emigrated to what later became impenetrable interested in stapeliads at one stage and Grootfontein, that any previously unrecorded Zimbabwe. There he set up a business supplying described several new species. He also published The stapeliads have been found since his time. Thus, for electrical equipment, especially batteries for vehicles. Stapelieae of Southern Africa (Lückhoff 1952) example, the extensive collecting done by Giess, His business was successful enough for him to sell it which, although it had only the briefest of texts, Merxmuller, Volk and Bleissner during the period off and 'retire' at the age of nearly 50. This gave him provided him with an opportunity to publish many of 1956-68, prior to the publication of Prodromus einer more time to pursue his programme of collecting and his remarkable black and white photographs. The Flora von Südwestafrika, revealed almost no new documenting the stapeliads and other succulents collecting activities between 1929 and 1956 of taxa of stapeliads. A few species not known to (mainly Euphorbia and Aloe) in Zimbabwe and A.G.J. (Hans Herre, who was really most interested Dinter were located by Ernst Julius Rusch and his other parts of south tropical Africa. He had begun in the succulent Aizoaceae and explored extensively son Ernst Franz Theodor Rusch. Both were farmers these studies in 1950 and continued them in in the less accessible parts of the Richtersveld, and businessmen who lived on the farm Lichtenstein Zimbabwe until he emigrated to South Africa in 1982. brought to light some new species. these were grown just south of Windhoek. They explored widely in the This research resulted in the publication of many at the Stellenbosh University Gardens and described by remote mountains of southern Namibia. papers on their taxonomy and culminated in his G.C. Nel, a botanist at the university, whose interests revision of Stapeliaa nd Huernia( Leach 1985; 1988). also lay more in the direction of the succulent In Moçambique, where stapeliads are Aizoaceae. comparatively rare, the only person who seems to Poth Pillans and Lückhoff lost interest in have taken a particular interest in them was the stapeliads later in their lives and it was only with the Portuguese agriculturalist Antonio de Figueiredo curatorship of M. Bruce Bayer at the Karoo Botanic Gomes e Sousa. He first saw stapeliads during a brief Garden in Worcester, from 1969 until 1987, that visit, in January and February of 1930, to the arid and interest came to be focused on them once more. A desolate Namibe (Mocamedes) district in southern programme of field exploration was undertaken and, Angola and was very enthusiastic when he found for the first time, populations were sampled several later that year in the Lebombo Mountains in extensively so that variation within populations Moçambique, a country where stapeliads were rather than just single plants could be observed. For virtually unknown. His interest in them continual the first time pollination and hybridization until 1947 and be wrote three accounts of the experiments were carried out and a comprehensive stapeliads of Moçambique (Gomes e Sousa 1935; herbarium record was initiated. LC Leach (see 1936; and the final one with Esteves de Sousa in below), who had already embarked upon a study of 1947). southern African stapeliads in preparation for an The natural history of Madagascar already began account for the Flora of Southern Africa project, to receive attention before 1658 (Reynolds 1966). also came to work at the Karoo Botanic Garden. He However, the southern parts of the island, which are stayed in this ideal environment from 1982 until home to most of its succulent species, including the 1990, greatly benefiting from the extensive collections stapeliads, were somewhat less accessible and so it of stapeliads that had been built up there. was only in 1918 that a stapeliad was first collected. Stapeliads were collected in Namibia as early as This was found by Raymond Decary and he 1878 by T.J.G. Een. Systematic exploration of the discovered a total of three species between 1918 and Namibian flora was begun by Moritz Kurt Dinter, 1932. The later exploration of the even more arid who was a horticulturist and botanist by training. At south-western corner of the island led to fur-there first he botanised alone and later often in the species being discovered by Montagnac and others. company of his wife, Jutta Dinter first arrived in In general, for individuals who have discovered Namibia in 1897 and left it for the last time in species, biographical information is given here only March 1935. He was especially interested in where no information is present in Gunn & Codd succulents and he expended much time and effort (1981). For further information the reader is referred exploring the arid south of the country where succulents to their book. are most common. He came across many stapeliads during the course of these travels. 3 Table 1. The subfamilies and tribes of the Apocynoideae (after Endress & Bruyns 2000) he stapeliads belong to the family Apocy-naceae, detailed and careful morphological studies over a areas and they become progressively rarer as one which is the seventh largest family of steadily increasing rage of taxa and from new moves away from these warmer part to temperate angiosperms and contains about 424 genera and molecular techniques by means of which parts of the regions. Most of them are herbs or climbers (trees some 4200 species. DNA of plants are analysed and compared. It are rare) and several of them have a weedy or When Linnaeus published his Species Plantarum indicates that the generally more derived Asclepiadaceae ruderal nature as is well known in the now almost of 1753, the species of Asclepiadaceae and has arisen within the Apocynoideae (e.g. Sennblad & cosmopolitan Arajia sericifera, Asclepias curassavica Apocynaceae were placed together in the large Bremer 1996). Several studies in which and Gomphocarpa fruticosus. The subfamily grouping which he termed 'Pentandria Digynia'. morphological and molecular data have been Asclepiadoideae is divided into four tribes This contained all those plants where each flower had analyzed together (e.g. Civeyrel et al. 1998; Asclepiadeae, Ceropegieae, Fockeeae and five anthers and two ovaries. In 1789 Antoine Sennblad et al. 1998) have shown that only a united Marsdenieae. The Fockeeae is sister to the other tree Laurent de Jussieu, who was one of the main Apocynaceae and Asclepiadaceae form a tribes, the Marsdeieae and Ceropegieae are sisters to proponents of Linnaeus's 'natural' system of phylogenetically acceptable 'monophyletic' unit. each other and together are sister to a Asclepiadeae. classification, took it a step further and placed all of Consequently, just under 200 years after Brown's The present account deals with the group of them in a 'family' (actually he called it an 'Ordo') important paper, it was proposed that the asclepiads know as to stapeliads and, in the past, which he called the Apocinae. This was named after the Apocynoideae and the Asclepiadoideae be united often referred to collectively as the Stapelieae. genus Apocynum, with which many of the species once more under a single family, the Apocynaceae Traditionally the tribe Stapelieae contained only the had at one time or another been associated. In 1810 (Endress & Bruyns 2000). The organization within highly succulent, practically leafless asclepiads with Robert Brown proposed that those genera in which this family is shown in Table 1. In this new angled stem (Brown 1902-03; 1907 09; White & the pollen was attached to ‘translators’ should be arrangement most genera of the former Sloane 1937) which, from the time of Linnaeus excised from the Apocynoideae and placed in a new Asclepiadaceae are placed within the subfamily (1753) up to that of Robert Brown (1810), were all family, the Asclepiadeae (Brown 1810). This is was Asclepiadoideae. The Asclepiadoideae, commonly, placed in the genus Stapelia. Closely related, less generally accepted and, since 1810, these two groups (though rather imprecisely) known as 'milkweeds', succulent taxa without angled stems were referred to were mostly treated most as separate families. includes some 2 700 species which are spread over 222 the tribe Ceropegieae (Brown 1902-03; 1907-09). Over the past three decades evidence has been genera. Representatives' of the Asclepiadoideae are However, Hooker (1885), who dealt with the accumulating from ever more most common in tropical to subtropical Rauvolfioideae Alstonieae 9 Old & New World, tropical to subtropical Vinceae 8 Old & New World, tropical to temperate Willughbeeae 18 Old & New World, tropical Tabernaemontaneae 19 Old & New World, tropical Melodineae 8 Old & New World, tropical to rarely subtropical Hunterieae 3 Old World, tropical Plumerieae 10 Old & New World, tropical to subtropical Carisseae 2 Old World, tropical to temperate Aiyxieae 7 Pacific Basin, Asia (1 genus in New World), tropical Apocynoideae Wrightieae 7 Old & New World, tropical to temperate Malouetieae 12 Old World (1 genus in New World), tropical to subtropical Apocyneae 27 Old & New World, tropical to temperate Mesechiteae 9 New World, tropical to subtropical Echiteae 22 Old & New World, tropical Periplocoideae 40 Old World, tropical to arid temperate Secamonoideae 9 Old World, tropical to temperate Asclepiadoideae Marsdenieae 29 Old & New World, mainly tropical to subtropical Ceropegieae 46 Old World, tropical to warm temperate includes the stapeliads, Ceropegia Asclepiadeae 145 Old & New World, tropical to temperate Fockeeae 2 Old World, tropical to warm temperate Number Subfamily Tribe of Distribution Commment genera 4 T Classification of the Stapeliads

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This important and long awaited two-volume monograph deals with the southern representatives of the fascinating group of succulent plants known as stapeliads. It is the only monograph on the topic published in the last 68 years. The book is the product of 25 years of research by the author who has b
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