Table Of ContentVolume 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
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Sponsors Edition: Vol. I - ISBN 1-919766-33-2. Vol. II - ISBN 1-919766-34-0 
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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
Description: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