TheNewy actus Jexicon Compiled and edited by David Hunt with the assistance of Nigel Taylor Graham (Charles and many members of the International (jactaceae Systematics Group Text dh 2006© Copyright 2006 David Hunt All rights reserved ISBN 0 9538134 4 4 (Complete Work) ISBN 0 9538134 5 2 (Text) ISBN 0 9538134 6 0 (Atlas) Published by dh books ‘The Manse Chapel Lane Milborne Port DT9 SDL England Printed in England by Remous Ltd Milborne Port SomersetThe New (actus Lexicon (Contents Contributors and Acknowledgments ........00000000c0eeeeeeee vi Early Subscribers Group Photographs ........0.0.e0e0ceeeeeeeeeeeeee weak Part I: Introduction Compiling the New Cactus Lexicon. . Notes on Classification Summary Table of Recognized Taxa Alphabetical Index of Genera Antificial Keys to Genera Conventions and Abbreviations Author Abbreviations ......... Book and Journal Abbreviations Selected Bibliography. . Part Il: Catalogue of Taxa...........- 25 Part III: Appendices 1. Recognized Taxa and their principal Synonyms . 283 321 322 324 V. Names in Current Usage 00.222. 0000ceeee eee 336 IL. Naturally-occurring Hybrids III. Unreferred Taxa IV. Conservation Assessments ..........vi (Contributors and Acknowledgments Contributors. Initial text drafts for this volume were compiled by the undersigned from original descriptions and other pub- lished sources, except those for the Brazilian Cereeae which ‘were compiled by Nigel Taylor from published treatments by or co-authored with his wife Daniela Zappi. The accounts cof various groups and genera were then revised in consultation with the specialist contributors listed here. Especial thanks for their input are due to Ralf Bauer (Hylo- cereeae and Rhipsalideae); Graham Charles (South American genera, especially Copiapoa, Espostoa, Gymnocalycium, Haageocereus), Martin Lowry (Echinopsis, Rebutia), Mats Hjertson (Echinopsis, Rebutia); Andreas Hofacker (Parodia); Reto Dicht & Adrian Luthy (Coryphantha); Detlev Metzing (Frailea, Gymnocalycium); John Pilbeam (Feracactus, Gym- nocalycium, Mammillaria, Rebutia) and Nigel Taylor (numer- ‘ous genera, especially Discocactus, Echinocereus, Escobaria, Ferocactus, Leptocereus, Rhipsalis and Stenocactus). ‘Advice and data, via publications, correspondence or attendance at ‘meetings has also been contributed by: Salvador Arias (Pereskiopsis ete), Geoff Bailey & John Miller (Ariocarpus), Wilhelm Barthlot (epiphytic cacti; comparative seed-morphology), Brian Bates (Rebutia etc), Pierre Braun (cacti of Brazil), John Brickwood (Parodia), Thomas Engel (Parodia), Urs Eggli & Daniela Zappi (compilers, Reper. Pl. Succ.), Roger Ferryman (Eriosyce), W.A. & Betty Fitz Maurice (Mammillaria), Ulises Guzmén (cacti of Mexico), Héctor Hernandez. & Carlos Gémez-Hinostrosa (Peniocereus etc), George S. Hinton (Geohintonia etc), Paul Hoxey (Turbinicarpus et), James Iliff (Andean Opuntioideae), Fred Kattermann (Eriosyce), Roberto Kiesling (Pierocactus ete), Myron Kimnach (Hylocereeac and Rhipsalideae), Paul Klaassen (Copiapoa), Wolfgang Krahn (cacti of Bolivia), Alfred Lau (Mammillaria), Beat Levenberger (Harrsia, Pereskia etc) Jonas Luthy (Mammillaria, Turbinicarpus), Jens Madsen (cacti of Ecuador), Massimo Meregalli (Gymnocalyci tum), Alessandro Mosco (Thelocactus), Roy Mottram (Cleistocactus, Copiapoa, Matucana), Reto Nyffeler (Austrocactus, Pfeiffera). Carlos Ostolaza (cacti of Peru), Donald Pinkava & Bruce Parfitt (North American Opuntioideae), Wolfgang Plein & Helmut Rogo- Zinski (Mammillaria), Gordon Rowley (nothogenera), Wolfgang ‘Stuppy (seed-morphology of Opuntioideae), Dieder Supthut (various genera), Baltasar Trujillo (cacti of Venezuela), Robert Wallace & Charles Butterworth (gene-sequence data). For a lst of contributors of illustrations, see the Atlas volume, p. vil. ‘Acknowledgments. Institutional support from successive Directors and senior staff of the Royal Botanic Gardens Kew enabled the compiler (D.H.) to participate in the activities of the International Organization for Succulent Plant Study (IOS) and the establishment in 1984 of the ad hoc IOS Cac- taceae Working Party, renamed in 2000 the International Cac- taceae Systematics Group, from which the present work and ‘others mentioned in the bibliography, have originated. The present Director of Kew, Professor Sir Peter Crane, is thanked for his positive encouragement for the project and for permit- ting Nigel Taylor, now Curator and Head of the Horticulture and Public Education Department, to attend these meetings and to continue making what has certainly been an indispen- sable contribution to its realization, Thanks are also due to Nigel Taylor, and to Graham Charles and Terry Palmer, for their help with proof-reading, and to the staff of Remous Ltd, Milborne Port, and especially Graham Bunter and Joe Fitzger- ald, for their care and patience during all stages of the pro- duction of both the Lexicon volumes. As publisher, 1 am further indebted to Ralf Bauer, Fred Kattermann and Dieder Supthut, who acted as treasurers in Germany, Switzerland and the USA during the pre-publication subscription period, and for the faith placed in the Lexicon project by all the ‘early sub- scribers’ listed on the following pages, who have helped to underwrite the cost of producing the work by subscribing up toa year in advance of its publication. Last but not least, I am much indebted to Karl Werner Beisel (Kakteenland Steinfeld, Germany) for his friendly support and ‘encouragement, and for lightening my task by providing me with many of the books and journals I have needed to consult; and to my wife Margaret Phillips, who might have believed, when I retired from Kew and we moved to the country, that I would forget about cacti and learn to play the organ properly. Instead, for the past decade and more, during her own busy career, she has found herself playing hostess to the frequent Visits of up to a dozen or more gentlemen with seemingly only ‘one topic of conversation, Such devotion speaks for itself! David Hunt Milborne Port I May 2006 ‘Atlas Volume Corrigenda Further o those listed om p. xiv ofthe Atlas volume, please note the following corrections tothe caption data 56.1 for *faseiculatus ssp. fasciculatus’ read engelmannit 613 for “E. gumeyiy’ read *(E. triglochidiatus var. gurneyi)” 98.4 and 99.1 for ‘speciosus’ read speciosus ssp. speciosus 202.4 for candelila read candelilla ssp. candelila 203.1 for candelilla read candelilla ssp. piray 227.4 for ‘Sardin de Cactus’ read Dinosaur Park 268.4 “insert data: BR, Rio Grande do Sul, Sd0 Francisco de Assis, HU 363 sis 2742 after “(21 insert (ssp. bayense) and replace data with: ‘AR, Buenos Arias, Olavarria, Sierra Baya, 180 m, Papsch 92.112-149, cult. (GC) 301.2 for ‘ssp. echinus’ read ssp. loccosa 4134 for “poselgeri read halei and for “W of La Paz’ read near San Carlos 481.2 for ‘moelleriana’ read moelleri 491.3 for ‘mesopotamica’ read megapotamica You can obtain an updated list of corrections by writing to dh books or by visting www.cactuslexicon.orgvii Early Subscribers ‘Anne Adams (UK) DrGR. Allcock (UK) Andrew (UK) Philip G. Andrews (UK) Per Andsager (Denmark) Eric Annal (UK) Dr Roberto Ariu (Italy) John L. Amold* (UK) Peter Arthurs (UK) ‘Torbjom Astlind (Sweden) Marco Avolio (Italy) Chris Backhouse (UK) Dr Geoff Bailey (UK) ‘Audrey & David Baines (UK) Dudley J. Baker (UK) Dorotiya Balizs (Hungary) Alberto Ballerio (Italy) Eddie Bamber (UK) Raiger Bamberg (Germany) Philip G. Barker® (UK) ‘Aymeric de Barmon (France) John Bamett (UK) Jamie Barron (UK) Dr Bob Barth (USA) Patrick Basset (Switzerland) Brian Bates (Bolivia) IT. 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Zimmerman (USA) Ben J.M. Zonneveld (Netherlands) Society, Library and other Corporate Subscribers Cactus & Succulent Society of Queensland (Australia) Cactus & Succulent Society of the ACT inc, (Australia) CCactusweelde afd, Tumhout (Belgium) Deutsche Kakteen-Gesellschaft* (Germany) DKG Echinocereus Group (Germany) DKG-Onsgruppe Offenburg (Germany) Kakteenfreunde Berlin e.V. (Germany) Massmann Internationale Buchhandlung (Germany) Succulenta (Netherlands) “Associacié d'Amics det Cactus *ASAC" (Spain) Schweizerische Kakteen-Gesellschaft (Switzerland) Sukkulenten-Sammlung Zirich (Switzerland) British Cactus & Succulent Society (UK) Bradford Branch BCSS (UK) Gloucester Branch BCSS (in memory of Rob Seward) (UK) Harrow Branch BCSS (UK) Liverpool Branch BCSS (UK) Manchester Branch BCSS* (UK) Northampton & Milton Keynes Branch BCSS (UK) ‘Nottingham Branch BCSS (UK) Portsmouth Branch BCSS (UK) Royal Botanic Gardens Kew (UK) Botanic Department, Chester Zoo (UK) Whitestone Gardens Lid* (UK) Gates Cactus & Succulent Society (USA) Michigan Cactus & Succulent Society (USA) San Gabriel Valley Cactus & Succulent Society (USA) Stockton Cactus & Succulent Society (USA) ‘Tucson Cactus & Succulent Society (USA) Gray Herbarium Library (USA) ‘The Huntington Library (USA) ‘The Luesther Mertz Library (USA) Arizona Cactus Sales Inc (USA) B&B Cactus Farm (USA) Rainbow Gardens Bookshop (USA) Cadillac Trading Company* (USA) ‘Twenty-one other subscribers indicated that they did not wish their name to be listed *Collector’s Editionx Group Photographs Botanists and musicians at a meeting of the IOS Cactaceae Working Party at the recently founded English Organ School & Museum (EOS), Milboe Port, April 1994, photographed by Roy Mottram. Left 1o right: Robert Wallace, Wolfgang Stuppy, Mary Iliff, Daniela Zappi, James Iliff, Nigel Taylor, Margarct Phillips, Ted(Compiling the New (actus Lexicon Much that Curt Backeberg wrote in the preface to his Kakteenlexikon (1966), translated into English a decade later by Lois Glass as Cactus Lexicon, would be apposite here. The purpose of the work is broadly the same and so, in general, is the format. In what was originally intended by Backeberg’s publisher to be a further edition of his work, we have preserved the basic elements: a summary of the classification adopted, an alphabetical listing of the taxa recognized, with concise descriptions and brief comments, and an extensive selection of illustrations (now in a separate volume, the *Atlas’). So in retain- ing the ttle Cactus Lexicon we are happy to acknowledge Backeberg’s unflagging indus- try in compiling his monumental six-volume handbook (Die Cactaceae, 1958-62) and his initiative in distilling it into that ‘handy work of reference’, as he called it, which has been the inspiration for ‘NCL! and one of its precursors. By the mid-1980s, when that new edition of the Kakteenlexikon was mooted, the virtual ‘monopoly on fieldwork and publication once enjoyed by collector-authors like Backe- berg and Friedrich Ritter was over. Ritter’s own prolix account of the cacti of southern South America, published in 1979-81, and the ofien conflicting views of the two collec- tors, would have made the editorial task of updating the original lexicon, with its supple- ment by Walther Haage, well-nigh impossible: in any case, more orthodox botanical treatments were becoming available for various individual countries with rich cactus floras, including Argentina, Brazil, Chile, Mexico and the USA; easier travel by air and on land was enabling many enthusiasts to fulfil their dreams of seeing cacti ‘in situ’ and to study them, their habitats and their variation, more critically; and the arrival of the per- sonal computer, the scanning electron microscope, gene-sequencing and other advanced laboratory techniques, was beginning to diminish the element of guesswork in the search for a generally acceptable classification at all levels Unlike Backeberg’s single-handed enterprise, therefore, the compilation of this New Cactus Lexicon has been very much a team effort, depending on the collaboration of many cactologists, some with formal training and professional experience in biological systematics but all it is fair to say, having started as amateur enthusiasts and growers. Most of us have had extensive field experience and between us we could probably claim to have seen almost all the recognized species in their native haunts. And whereas Backe- berg was constrained to illustrate just one or a few species in each genus he recognized, ‘we have attempted to illustrate ali the taxa we can currently recognize and to do so with pictures of plants in habitat or of documented origin. We have not quite succeeded but believe that the Atlas volume, with its selection of just over 2500 of the many thousands of images submitted, is the best-documented photographic record of the diversity of the ‘cactus family yet published. For a group of plants described by Richard Bradley in his History of Succulent Plants (1716-27) as ‘incapable of an hortus siccus’ and, to this day. very inadequately represented by preserved material in the world’s herbaria, the illustra- tions can serve as a kind of embyronic pictorial herbarium. Though they are only virtual specimens, ‘incapable’, one might say, ‘of forensic examination’, they and their labels (the captions) are certainly capable of discussion, confirmation or revision, of encourag- ing us to look more closely at the plants in our collections and the original descriptions, and of tempting us to pack our bags and cameras and head off to that El! Dorado which Backeberg called Stachlige Wildnis, the prickly wildemess. ‘The catalogue of taxa, which forms the main part of the text, has largely evolved by a process of updating and elaborating previous compilations, most recently the second edition of the CITES Cactaceae Checklist (1999), in itself a mini-lexicon, as is any ‘checklist’, being. in the words of one famous example, The Shorter Oxford Dictionary. ‘a word-book or dictionary; a special vocabulary; a lst of words or names” — but not (and the point is an important one!) a monograph or an encyclopaedia. Like Backeberg's, ours, is essentially a dictionary of the genera, species and subspecies of cacti recognized by our principal contributors on the basis of their current knowledge of the plants inthe field and cultivation, and of the literature. It does not include chapters of general information about the family, its history, morphology, cultivation and so on, such as may be found in several other treatises, and makes no pretence to be monographic or encyclopaedic in2 Compiling the New Cactus Lexicon scope or to present a definitive or fully worked-out classifica tion By way of background, a Working Party was first convened in 1984 under the auspices of the International Organization for Succulent Plant Study (IOS) and has met regularly ever since, changing its name in 2000, at the suggestion of some of its members, to the International Cactaceae Systematics Group. ‘An initial ‘consensus’ list of genera accepted by the partici- pants (Bradleya 4: 65-78. 1986) was intended to serve as ‘a framework for continuing discussion’ and soon prompted the Nomenclature Committee of CITES (the Convention on Inter- national Trade in Endangered Species of Fauna & Flora) to ‘commission the preparation of the first edition of the CITES Cactaceae Checklist (1992). The generic ‘consensus’ lst also formed the framework for treatments of the Cactaceae by Hunt & Taylor for The European Garden Flora (Walters etal 1989) and by Barthlott & Hunt for The Families and Genera of Vascular Plants (Kubitzki et al., vol. 2. 1993) and the basis ‘of various research topics, leading in their turn to changes in the classification adopted in the second edition of the Check- list and yet more in the present work. Basic name-list As a prerequisite for the Checklist, abasic but comprehensive electronic name-list of over 12,000 names of proposed cactus species. with their authors and literature ref- erences, was compiled from several sources. The names were “status-coded” to indicate (a) whether they were basionyms the names applied to taxa when first described) or classi- ficatory changes (‘combinations’, transfers to a different genus or changes of rank, such as species to subspecies); and (b) whether or not they had been used as an accepted name anywhere in the ‘benchmark bibliography’ of mainly recent or standard literature and could thus be considered to be ‘in current usage’. Transfers and changes of rank were linked to the relevant basionym by way of the record number of the basionym. For the second edition of the Checklist names at subspecific rank were added to the name-list. For both edi tions and the present work, the deadwood of names not ‘in ‘current usage’ has generally been filtered out, leaving a net total listed in this work of c. 7700 names. Format of entries in the Lexicon. Entries for recognized taxa normally consist of two or three short paragraphs, the first containing the name, source and distribution data, the second the description, and the third any relevant comments. Entries for taxa included in the catalogue but not currently recognized may also consist of two or three paragraphs, but more com- monly a single paragraph indicating the taxon to which they are referred. The standard sequence of data in the first and second paragraphs is explained on p. 13. Author and literature references. The tradition of appending the name of the author or authors to the formal citation of botanical name grew up at a time when authors and their books were few, and the modem rules of nomenclature ‘unwritten. Today, mere citation of authorship (unless the cita- tion refers to a name used in an incorrect sense) may convey an impression of authority, but is of litle practical use without a reference to the place and date of publication as well. The brief for the CITES Cactaceae Checklist did not include author-citations, but as the compiler received numerous requests from individual users of the first edition to include them in the next, an abbreviated format was devised to enable each name and its reference to be printed on a single line. In the event this format was not used, but it received a favourable response when used in the Working Party's bulletin and, having proved very convenient editorially, has been adopted here. It is explained on p. 13. Basionyms and nomenclatural type data. As well as giving author and literature references, it has been the policy in com- piling the Lexicon entries to indicate, where practicable, the nomenclatural type of each accepted name, preceded where applicable by its basionym (the original name on which a name used in a later classification is based). Rules have been introduced over the years to ensure that the precise meaning or application of names newly proposed is clear from the ‘outset (or, for older names, can be fixed later) by reference to an individual preserved specimen, or if necessary an illustra- tion, termed the ‘1ype’. The type (which does not have to be an average or ‘typical’ specimen of the species or other taxon concerned) gives the botanist an absolutely fixed point of ref- ‘erence from which to judge whether other specimens to which the name has been applied are correctly identified or not. For this reason, a type specimen should be as complete as possi- ble, with flowers and or fruits and seeds, as well as vegetative material, so that its identity will be clear and unambiguous. Sadly, Backeberg’s failure to preserve material of the names he proposed was notorious and the scrappy material deposited in herbaria by Ritter and others to satisfy the letter of modem rules, though not their intention, has greatly exacerbated the Jifficulties of those working on the taxonomy of the family. ‘Synonyms. In general, the Catalogue of Taxa does not include names considered for our purpose to be ‘in current usage’ if they are not the accepted name of a recognized taxon or its basionym. This is parlly to avoid cluttering what is primarily ‘an annotated list of taxa we accept with names we do not, but also because it has been our policy not to amplify the descrip- tions of taxa we recognize to cover other taxa treated ~ often only provisionally or tentatively - as ‘synonyms’ (‘subsumed taxa’ would be a better term), except in the genera for which up-to-date botanical revisions exist (those with “A’ or “B” status in the Table on p. 5). In genera for which such revisions do not exist, separate entries or partial entries are often pro- vided in the main catalogue for the subsumed taxa, and an alphabetical list of synonyms in current usage occupies the final pages of the volume (Appendix V, p. 336 et seq,), along with the accepted names. A summary of recognized taxa with their principal putative synonyms is given in Appendix I. Further synonyms are accounted for in the main catalogue if they are mentioned in a caption in the Atlas, if more familiar than the current name (such as those of the many species of Opuntia now included in separate genera not recognized in the published editions of the CITES Cactaceae Checklist), or if previously accepted or provisionally accepted in the Checklist ‘but here subsumed under another species. Naturally-occurring, hybrids mentioned in the main text are also listed in Appendix I, and some names which are neither accepted nor referred to «a recognized taxon, because they are problematical (prefixed °@ in the main text) or else because insufficiently or not yet studied, will be found in Appendix IlNotes on Classification 3 Geographical distribution. As yet, reliable and detailed distri- bution data are available for a relatively small proportion of the recognized taxa. So, as a minimum, we indicate the country or countries whence they have been reported, using ISO codes (see p. 13), and for the larger countries the names of the primary political divisions (states, departments or provinces etc). Further habitat information including vegeta- tion types and altitudinal ranges may occasionally be given if ithas been provided by contributors, but systematic coverage has been beyond the scope ofthis compilation. Descriptions. The concise format devised for these by Backe- berg, mentioning the principal features of the vegetative and reproductive parts, with each phrase preceded by a bold-type abbreviation for the relevant part, is used here (for the abbre- viations used, see p. 13). In principle, the aim has been to confine the text to the diagnostic features of the taxon con- cemed, derived either from a reliable modem source, or else the original description. Intermediate sources, which may not be authentic or reliable, have been avoided. It is necessary, however, to make the caveat that some of the descriptions may be difficult to reconcile with the supporting illustrations in the atlas volume, especially in genera where there is as yet no reliable modem source. Notes on (lassification Bill Bryson, in his Short History of Nearly Everything, writes Faxonomy is sometimes described as a science and some- times as an art, but really it’s a battleground.” Taxonomists would probably prefer to say their subject is a meeting-point of science and art, involving the application of scientific rigour to the development of a system of classes or taxa within an artificial hierarchy and an agreed set of ground- rules. It is a meeting-point that allows ample scope for philo- sophical debate and for the comparison and evaluation of different hypotheses in the light ofthe available data but also, and here is the crunch, for the names of the taxa to be differ- ent according to the classification preferred by different users ‘or adopted in works such as this. Above the rank of genus this concems ordinary users very little, and though the advent of molecular systematics is now having a significant impact on the taxonomy of the family Cactaceae at the rank of subfami- |p, tribe and subtribe, the names of the taxa at the equivalent ranks have remained relatively stable for more than a century. Subdivisions within genera, above the rank of species, are also fairly uncontroversial, since they do not affect the plant- names in everyday use. It is only at the ranks of genus and species that cactus taxonomy has shown the capacity to degenerate into a battleground, a kind of ‘class’ war, one right say, between the so-called ‘lumpers’ and ‘splitters’, leading to what might be regarded as another kind of ‘prickly wilderness’. Subfamilies and Tribes. Whereas the Lexicon text is arranged alphabetically by genus etc, the Atlas illustrations are grouped under the relevant subfamilies and tribes, and with the genera in a sequence partly pragmatic and partly guided by current views on their relationships (see Atlas p. ix-xi). Three sub- families were proposed by Schumann in 1898, Cereoideae (now Cactoideae), further divided into three tribes with a total of 16 genera, Pereskioideae (Maihuenia and Pereskia) and Opuntioideae (3 genera). In the light of initial molecular data a fourth, Maihuenioideae (Mailuenia), was proposed in 1996. Meanwhile, Britton & Rose (1919-23) treated Schumann's three subfamilies as tribes, placing Maihuenia in the Opun- tieae and dividing the Cereeae into eight subtribes. Until recently this division of the equivalent subfamily Cactoideae. in a re-working with nine tribes by Buxbaum (1958), was widely followed, but molecular data now point to there being fewer major clades in the family, suggesting the expansion of tribe Echinocereeae to include the other cereoid groups with spiniferous flowers and fruits (tribes Leptocereeae and Pachycereeae). Molecular studies also suggest that subfam ‘Opuntioideae could be divided into into five tribes, but, as the nomenclature of these has not yet been formalized, a simpler arrangement in two tribes, Cylindropuntieae and Opuntieae has here been adopted for the illustrations in the Atlas. Genera. It is here, since Britton & Rose chopped the 21 genera recognized by Schumann into 124, that there could indeed be said to have been a battleground in cactus taxono- ‘my during the twentieth century. While the genus is an inher- ently more subjective concept than the species, and in theory there is nothing wrong with preferring lots of litle genera to the old system with fewer, larger ones, the Britton & Rose approach has led to enormous instability in the names of the plants, by greatly increasing the likelihood of individual species being moved from one small genus to another, for some of the smaller genera to be recombined, and so on. What Britton & Rose did to the Cactaceae and other groups they fragmented was condemned by other taxonomists as “cactusization’ and by the more recent monographer of the North American cacti, the late Lyman Benson, as ‘the height of influence of a local liberal school in the United States’ which ‘flourished from about 1900 to 1930" and whose ‘poli- cies have been abandoned or modified by the great majority of later botanists’, In Europe, among cactologists, these included Alwyn Berger and Schumann’s successor, Friedrich Vaupel but not, of course, Backeberg, Buxbaum and Ritter, who between them further doubled Britton & Rose’s generic total to around 250. Backeberg’s own final tally, in the Kakteen. Texikon, was 233, By the 1980s the ‘war’ between Backeberg (who had died in 1966 just before the Kakteenlexikon was published) and the others, coupled with their incompetence in matters nomen clatural and Backeberg’s self-confessed ‘partial reserve to some antiquated rule-details’, had brought about a situation verging on anarchy where, in the worst cases, a single species could have synonyms in half-a-dozen or more genera. With the passing of the main combatants, however, a more concil- iatory attitude has developed, and also, since the advent of ‘molecular systematics, a greater willingness to ‘wait and see’ before making further changes at generic level. Since the IOS Cactaceae Working Party published its first report in 1986, a degree of stability can be said to have retumed. Several changes have, however, been agreed since 1999 and the second edition of the CITES Cactaceae Checklist (108 genera accepted, excluding nothogenera). The main change (some would say reversal) since then, has been the re-division of4 Notes on Classification Opuntia, adding 12 genera. Three small genera provisionally accepted in the Checklist are not accepted here (Acanthocaly- cium, Cephalocleistocactus and Cintia), but the recognition (for the time being, atleast) of seven small genera on the basis ‘of molecular o other evidence (Acharagma, Castellanosia, Cumarinia, Pfeiffera, Strophocactus) or else newly described, (Pierrebraunia, Yavia), has brought us back, in this work at least, to 124 genera, coincidentally the same as in Britton & Rose's monograph. Species. Given that this is generally understood to be the basic unit of classification, one might expect it to be capable of a reasonably straightforward definition. But it isn’t, and more so for plants, which as individuals are static, than for animals, ‘which can mostly run, swim or fly. One can simply avoid the issue and admit all-comers by saying a species is ‘one of the categories of taxa recognized in the Intemational Code of Botanical Nomenclature, that to which a binomial is given’: ‘or, as we do here, more helpfully perhaps, ‘a series of similar intergrading and interfertile populations, recognizably dis- tinct from other such series and reproductively isolated from other such series’. In theory this basic definition covers pop- ulations which are capable, genetically, of interbreeding but do not do so in nature because they are remote from one another or effectively isolated by other biogeographical or ecological barriers. AS a species becomes established over an appreciable range, or in separate populations, however, itis to bbe expected that its individuals will only interbreed locally ‘and gradually accumulate consistent differences from those in ‘other populations, and the more so if the plants are self-fertile. When the differences are maintained in cultivation, as is often the case with the smaller cacti popular with collectors, and appear to have a genetic basis, the question certainly arises as {to what is meant by ‘recognizably distinct’ and the answer will fofien depend on a personal point of view or special interest. For anyone growing or collecting living plants, the desire for the names to be precise and constant is paramount. In essence, and justifiably, the approach should be that of the nursery- ‘man’s catalogue so as to be able to specify exactly the plant fone has, or is looking for, or is talking about. For every item that looks different or behaves differently, a different name, and the simpler the better, is what is needed. For taxonomists, however, the emphasis is on classification For them the species is a more elastic concept, since itis sup- posed to take into account the variability within and between populations, a point often neglected by many collector- authors in the past. For the most part, the cultivated specimens derived from their introductions have either been raised from seed, originally collected from just one or a few wild individ- uals, or else propagated from offsets etc, and in fact ‘cloned’ from one individual specimen or just a few. What we have in ‘our collections are thus ‘selections’ (what one distinguished taxonomist called “flowerpot species’), and that they seem ‘more or less constant, invariable or predictable, and distinct, is not very surprising. But it doesn’t mean they are all different species in the sense defined above. Enthusiasts who have Visited the habitats realise that many species are much more variable out there than is apparent from the plants in their col- lections and that the best way to specify exactly the plant one is talking about is to relate it to its precise source, via an indi- vidual locality-name or a collector's field number. The field- work and publications of this new generation of enlightened “cactus explorers’ are greatly helping to correct the very narrow concept of species that was promoted by Backeberg, Ritter and others. Subspecies. Much-needed rationalization of species-concepts in popular groups of cacti has also been helped in the past decade or s0 by the use of the category of subspecies for sig- nificant variants, especially those which represent groups of Populations occupying more or less discrete areas within the overall range of a species. This has followed moves by the IOS Working Party to simplify the nomenclature of the ‘Iess- than-species’ and to recommend the use of subspecies instead of the more familiar but ambiguous category of variety, long used indiscriminately for both naturally-occurring and horti- cultural variants (now preferably treated as cultivars). In this ‘work subspecies is the only infraspecific rank formally recog- nized, except where the basionym of an accepted name was published at varietal (var.) or forma (f.) rank. All trinomials listed in the Appendices are thus intended at subspecific rank, and the rank indicator (or ‘connecting term’) “ssp.” given in the more formal Catalogue entries is dispensed with. Summary Table of Recognized taxa. The table opposite lists the genera (other than hybrid genera) recognized in this work, and the number of taxa (species and (+) heterotypic sub- species and total) recognized in each of them. For comparison, the total number of taxa per genus recognized in the second edition of the CITES Cactaceae Checklist (1999; table on pp. 159-160), excluding hybrids (10), is also given (CCC2) together with the net difference (+/-), ‘The overall net reduction (434) in the number of taxa recog- nized here (many former Opuntia species now being referred to the segregate genera, of course) represents approximately 19% of the previous total. Conservative floristic treatments, such as those in the Flora of North America and Cacti of Eastern Brazil, and specimen-based revisions such as that of the tribe Hylocereeae by Ralf Bauer, account for part of this reduction, and the rest largely result from examination of the {650+ species and heterotypic subspecies provisionally accept- ed in CCC2, many of which, being insufficently known or “data deficient’, are not now considered to be ‘recognizable’, at least until their status has been clarified by specimen-based research and fieldwork. Where practicable they have been referred on a ‘best guess’ basis to a related species, but might better be regarded as ‘subsumed taxa’ than as “synonyms”. ‘The letters A~D in the ‘Status’ column in the table are intend- ed to give an indication of the relative reliability of the taxon- ‘omy adopted here for the genera (82) with more than two recognized taxa, based mainly on the principal source(s) : ‘A. Monograph (Pereskia) B_ Regional or partial specimen-based revision ete (27) C__Semi-popular handbook/study-group publication(s) and/or older regional treatments (37) D_ Provisional assessments, often protologue-based (17)Summary Table of Recognized Taxa 5 Genus Taxa | Total CCC +/- [status] Genus Taxa | Total CCC2] +/ [status ‘Acanthocalycium 3} af — Leuchtenbergia reo] 1] a] o ‘Acanthocereus r+0] 1] 6] s]d 340] 3} 2] 1] c ‘Acharagma 241] 3] 0] 3 2+0] 2} 2] ol a ‘Ariocarpus 7+1] 38] 8] of B 7+0] 7] 0] 7] ¢ ‘Armatocereus 742] 9] 15] «| D Mammillaria 163+ 69 | 232 | 264 | 32 | Arrojadoa s+o] s] 6] +] B Mammilloydia t+o}] i] a] oo ‘Arthrocereus 4+2] 6] 6] o| B Matucana we7]afa} ofc ‘Astrophytum 6+0} 6} 4] 2] ¢ Melocactus 37+ 13. | 50] 45 | 5] B ‘Austrocactus 340] 3] s]| 2} c¢ Micranthocereus 9+i} | 9} i] B Austrocylindropunia} 8+ 1 | 9 | 0] 9] © Mila 142] 3] 4] 2] oD ‘Aztekium 20] 2] 2] 0 Miqueliopuntia reo] i] o] t Bergerocactus r+o} a] a] o Myrtillocactus 4+0] 4] 4] ofc Blossfeldia t+of ir} i} o Neobuxbaumia aso} ae] 9} alec Brachycereus r+o} 1} 1} o Neolloydia 240] 2] 2] 0 Brasilcereus 2+0] 2} 2} 0 Neoraimondia 240] 2) 3) 4 Brasiliopuntia r+o} i} of] 4 Neowerdermannia | 2+0] 2] 3 | -1 Browningia g+if} 9} uu} 2] od Nopalea 440] 4] 0} 4] od Calymmanthium reo} 1} 1} o [Obregonia treo} a] afio Camegiea reo} i} i} o Opuntia 75+ 0 | 75 | 361 |-2%6 | b Castellanosia t+o} 1} of] a /Oreocereus 6+2]/ a8] 9} alc Cephalocereus 340] 3] s} 2] ec JOroya 240] 2) 2] 0 Cephalocleistocactus r} al — JOnegocactus reo} ail] a] o Cereus +4] 29 | 39 | 10] dD Pachycereus Beolpfea] afc Cinta ry als Parodia sg +8 | 66 | 99 |-23| dD Cipocereus s+1i] 6} 6] o| B Pediocactus 7+3] 0] 6 | 6] c Cleistocactus 38 +10 | a8 | 59 | -] dD Pelecyphora 2+0/ 2) 2] 0 Coleocephalocereus | 8+ 2] 10 | 8 | 2| B Peniocereus 2+0] 20] 8] 2\| c Consolea 340] 3] 0} 3] D Pereskia +t] is} is} ala Copiapoa a+9o] 3] 29] if c Pereskiopsis 640] 6| 7] 1] B Comyocactus +0] 12 | 35 | 23] D Preitfera 941] 0} 0} wo] c Corynopuntia +o] ia] o] wi c Pierrebraunia 240] 2] 0} 2 Coryphantha aeu] 3] sr} 4] c Pilosocereus 4+] 49] a3 | 6] B Cumarinia reo} ar] of] 4 Polaskia 240] 2] 2] 0 Cumulopuntia 4+3/ 7] 0] 7] c¢ Praecereus 244] 6] 6} o| c Cylindropuntia 3+o | 33] 0] 33] B Pseudoacanthocereus | 2+ 0 | 2 | 2 | 0 Dendrocereus 240] 2] 2] 0 Pseudorhipsalis 6+3] 9] 6] 3] B Denmoza t+o}] a] a] o Prerocactus 9+0] 9] 9] o] B Discocactus u+i1{2]o] 2] ¢ Pygmacocereus 240] 2] 3] -1 Disocactus uss |||] o] 6 Quiabentia 240] 2] 2] 0 Echinocactus 6+1]/ 7] 7] o] B Rauhocereus r+1] 2] 2] 0 Echinocereus 67 +39] 106 Ji | os] B Rebutia 2+ | | 53 |-3] Dd Echinopsis 77+ 2% | 101 | 149 | 48 | D Rhipsalis 35413 | 48 | 48 | o| B Ej 2+6] i] is] a} B Samaipaticereus iso] 1] a] oo Epithelantha 2+0/ 2] 6] 4] B Schlumbergera 6-0] 6] 7] 41] 8 Eriosyce s2+19] si | | 3] ¢ Sclerocactus 20+ 4] 24] 2 | a] c Escobaria 944] 23] 30} 2] ¢ Selenicereus +3] 1s | 7 |-2] B Escontria treo} a} a} oo Stenocactus +o] 8| wo | 2] c Espostoa nei] 2] wl 4a} c Stenocereus wei] 2] | of ¢ Espostoopsis reo} a] a} oo Stephanocereus 240] 2] 2] 0 Eulychnia 44a] 5} 7] 2] ¢ Stetsonia treo} i] i] oo Facheiroa 341] 4] 4] o] B Strombocactus rer] 2] 2] 0 Ferocactus we] a2] 39] 3] c Strophocactus 3+0] 3] 0] 3] 8 Frailea +6] is | 2] 7] D Tacinga o6+2/ 8] 2] 6] B Geohintonia reo} rf a} o Tephrocactus 740] 7] of 7] ¢ Grusonia +o} 1} of] a ‘Thelocactus w+6] 2] a} a] c Gymnocalycium 49414] 6 | 90 | 27] ¢ Tunilla s+o] s] of} s] po Haageocereus 945] 14] 25 | -1} D Turbinicarpus +20] 3% | 3 | 3) c Ham 940] 9] 2} -2] d Ucbelmannia 343] 6] s| 1] 8 Hatiora 6+i] 7] 6] afc Weberbauerocereus | 7+ 0] 7] 8 | a] Cc Hylocereus +o] a] is] 4a] B Weberocereus +2] 0] 9] 1] B Jasminocereus reo] 1] a] o Yavia 140} rf oo] 4 Lasiocereus 240] 2] 2] 0 ‘Yungasocereus reo} 1] a] o Leooereus 1+0] 1] 1] of B Lepismium 6+0] 6] is} 9] c . eet Mba cal aed elec Totals 1438 +378| 1816 | 2250 | -4346 Index of Genera Alphabetical Index of Genera Numbers before the @ symbol refer to the relevant pages in this volume, and those after the symbol fo those in the Atlas. Acanthocereus .. =. 25 20 Acanthocalycium —+ Echinopsis Acantholobivia —> Echinopsis Acanthorhipsalis B+R. —> Peiffera Acanthorhipsalis Kimn —> Pfeiffera Acharagma .. -26 & 393 Akersia —> Cleistocactus Ancistrocactus —» Sclerocactus Anisocereus > Pachycereus Aporocactus > Disocactus Arequipa —> Oreocereus Arequipiopsis > Oreocereus Ariocarpus = 26 @ 363 ‘Armatocereus 27@ 12 Arrojadoa . 28 @ 158 Arthrocereus = 30@ 229 Astrophytum = 31 338 Austrocactus ..... 3108 Austrocylindropunti 32 @ 456 Aztekium 34.@ 340 Azureocereus > Browningia Backebergia > Pachycereus Bartschella —> Mammillaria Bergerocactus 34027 Binghamia —> Espostoa Bisnaga —> Ferocactus Blossfeldia 34.327 Bolivicereus > Cleistocactus Bonifazia > Disocactus Borzicactella —> Cleistocactus Borzicactus > Cleistocactus Brachycalycium —> Gymnocalycium Brachycereus - 34. 180 Brasilicactus —> Parodia Brasilicereus 34@ 137 Brasiliparodia —> Parodia Bravocactus > Turbinicarpus Brasiliopuntia 35 @ 485 Browningia 35 @ 124 Buiningia -> Coleocephalocereus Cactus L > Mammiltaria Cactus sensu B+R —> Melocactus Calymmanthium ..........36@ 1 Carnegiea.... -36 34 Castellanosia 2376 Cephalocereus 37039 Cephalocleistocactus > Cleistocactus Cereus 37 @ 129 Chamaecereus > Echinopsis Chiapasia > Disocactus Chichipia + Polaskia Chilita > Mammillaria Cintia + Rebutia Cipocereus . 42 @ 137 Cleistocactus =. 43 @ 201 Cochemiea —> Mammillaria Cochiseia > Escobaria Coleocephalocereus ..... 49 & 160 Coloradoa —> Sclerocactus Consolea 51 @ 485 Copiapoa 52 € 328 Corryocactus 5685 Corynopuntia 60 @ 479 Coryphantha 62 @ 381 Cryptocereus > Selenicereus Cullmannia —> Peniocereus Cumarinia 67 & 393 ‘Cumulopuntia 67 @ 459 Cylindropuntia 69 & 470 Deamia — Strophocactus Delaetia + Eriosyce Dendrocereus =. 74220 Denmoza 748214 Digitorebutia > Rebuia Digitostigma —> Astrophytum Discocactus 74 @ 287 Disocactus 11095 Dolichothele > Mammillaria Eccremocactus > Weberocereus Echinocactus =. 79 336 Echinocereus . - 80 0 54 Echinofossulocactus Lawr — Echinocactus Echinofossulocactus sensu B+R > Stenocactus} Echinomastus > Sclerocactus Echinopsis - 90 231 Emorycactus —> Echinocactus Encephalocarpus —> Pelecyphora Eomatucana —> Matucana Epiphyllanthus > Schlumbergera Epiphyllopsis -» Hatiora Epiphyllum . =. 104 @ 90 Epiphyllum Pf > Schlumbergera Epithelantha 106 & 36 Endisia + Corryocactus Eriocactus —> Parodia Eriocephala > Parodia Eriocereus —> Harrisia Eriosyce 106 @ 292 Enythrorhipsalis —> Rhipsalis Escobaria 112 @ 394 Escobariopsis > Mammillaria Escobrittonia > Coryphantha Escocoryphantha —> Escobaria Escontria 15 @41 Espostoa 115 @ 188 Espostoopsis 117 @ 156 Eulychnia . 117 @ 10 Facheiroa .. 118 @ 193 Ferocactus 118 & 368 Floribunda > Cipocereus Frailea 122 @ 324 Geohintonia = 125 @ 340 Gerocephalus — Espostoopsis Glandulicactus —> Sclerocactus Grusonia 126 @ 481 Gymnanthocereus > Browningia Gymnocactus — Tutbinicarpus Gymnocalycium 126 & 268 Haageocereus 135 @ 183 Hamatocactus —+ Thelocactus Harrisia 137 @ 26 Haseltonia + Cephalocereus Hatiora 138 @ 120 Hylocereus 139 @ 85, Hymenorebutia —> Echinopsis Islaya > Eriosyce Isolatocereus > Stenocereus Jasminocereus .. 141 @ 15, Kadenicarpus —» Turbinicarpus Krainzia — Mammillaria Lasiocereus - 141 @ 195 Lemaireocereus > Pachycereus Leocereus 141 @ 180 Lepidocoryphantha > Coryphantha Lepismium 142. @ 104 Leptocereus 143 @ 17 Lepiocladodia > Marnmillaria Leuchtenbergia 144 @ 367 Leucostele —> Echinopsis Lobeira > Disocactus Lobivia > Echinopsis Lophocereus > Pachycereus Lophophora . . = 144 @ 362Index of Genera 7 Lowanthocereus — Cleistocactus Lymanbensonia —> Preiffera Machaerocereus —> Stenocereus Maihuenia .. 145 @ 455 Maihueniopsis ......... 145 @ 461 Malacocarpus S-D non Fisch+Meyer > Parodia ‘Mamillopsis > Marnmillaria Mammillaria 147 @ 400 Mammilloyd 180 @ 367 Marenopuntia > Opuntia Marginatocereus > Pachycereus Maritimocereus —> Cleistocactus Marniera > Selenicereus Marshallocereus —> Stenocereus Matueana 181 @ 218, Mediocacuus > Hylocereus Mediolobivia Bkbg > Rebu Melocactus 183 @ 164 Meyerocactus > Echinocactus Micranthocereus ....... 190 € 153 Mila ooo. c ecco. 19 @ 181 Miqueliopuntia 192 @ 482 Mirabella > Cereus Mitrocereus —> Pachycereus Monvillea B+R (as to type) > Acanthocereus Monvillea sensu B+R —> Praecereus Morangaya > Echinocereus Morawetzia > Oreocereus Myrtillocactus 192 @41 Navajoa > Pediocactus Neoabbottia > Leptocereus Neobesseya —> Escobaria Neobinghamia > xHaagespostoa Neobuxbaumi 192 @ 35 Neocardenasia > Neoraimondia Neochilenia > Eriosyce Neodawsonia -> Cephalocereus Neoevansia -> Peniocereus Neogomesia —> Ariocarpus Neolloydia .. 193 @ 365 Neomammillaria > Mammillaria Neoporteria —> Eriosyce Neoraimondia . 194 @ 15 Neowerdermannia ..... 194 @ 291 Nopalea ... 195 @ 487 Nopalrochia —> Disocactus Normanbokea —> Turbinicarpus Notocactus —> Parodia Nyctocereus -> Peniocereus Obregonia 195 @ 362 Ochmea >» Mammillaria Opuntia + 196 @ 488 Oreocereus . ce 214 @ 214 Oroya oe. cee es 215 225 Ortegocactus. 216 @ 399 Pachycereus 216 @ 27 Parodia : -217 & 306 Pediocactus. .. 225 @ 341 Pelecyphora -226 @ 399 Peniocereus. 226 @ 24 Pereskia. 229 @ 450 Pereskiopsis coos 231 © 468, Peruvocereus > Haageocereus Preiffera . 2312 Phellosperma —» Mammillaria Philippicereus —» Eulychnia Phyllocactus > Epiphyllum Pierrebrauni 233 @ 156 Pilocanthus > Pediocactus Pilocereus Lem > Cephalocereus Pilocereus KSch > Pilosocereus Pilocopiapoa > Copiapoa Pilosocereus 233 @ 139 Polaskia. 241 @ 43 Praecereus coves 241 128 Pseudoacanthocereus. .... 242 @ 21 Pseudoespostoa —> Espostoa Pseudolobivia > Echinopsis Pseudomammillaria —» Mammillaria Pseudomitrocereus > Pachycereus Pseudonopalxochia > Disocactus Pseudopilocereus > Pilosocereus Pseudorhipsalis 242 @ 101 Pseudozygocactus > Hatiora Plerocactus............243 € 466 Plerocereus > Pachycereus Puna —> Maihueniopsis Pygmaeocereus.........244 € 182 Pyrrhocactus —> Eriosyce Quiabentia 244 © 468 Rapicactus —> Turbinicarpus Rathbunia —> Stenocereus Rauhocereus 245 @ 196 Rebutia = 245 @ 256 Reicheocactus -> Echinopsis Rhipsalidopsis —> Hatiora Rhipsalis Rhodocactus > Pereskia Ritterocereus > Stenocereus Rodentiophila > Eriosyce Rooksbya > Neobuxbaumia Roseocactus > Ariocarpus Roseocereus > Harrisia = 253 106 ‘Samaipaticereus 257 & 226 Schlumbergera «258 @ 122 Sclerocactus 259 @ 343 Selenicereus 262 @ 82 Seticereus —> Cleistocactus Seticleistocactus —> Cleistocactus Setiechinopsis > Echinopsis, Siccobaccatus > Micranthocereus Soehrensia —> Echinopsis Solisia > Mammillaria Stenocactus 264 € 379 Stenocereus. 265.0 44 Stephanocereus 268 @ 157 Stetsonia .. 268 @ 127 Strombocactus 268 © 361 Strophocactus. 268 @ 22, Submatucana —> Matucana Subpilocereus —> Cereus Sulcorebutia > Rebutia Tacinga 269 & 483 ‘Tephrocactus . = 270 464 Thelocactus. .. 271 349 Thelocephala > Eriosyce Thrixanthocereus —» Espostoa Torreyocactus > Thelocactus Toumeya —> Sclerocactus Trichocereus > Echinopsis, Tanita. 273 @ 482 ‘Turbinicarpus. 274 @ 354 Uebelmannit 278 @ 178 Utahia > Pediocactus Vatricania > Espostoa Weberbauerocereus..... 278 @ 197 Weberocereus 2.279 80, Weingartia > Rebutia Werckleocereus > Weberocereus Wigginsia —> Parodia Wilcoxia — Echinocereus Wilmattea > Hylocereus Winteria Ritt non Murr ~ Cleistocactus Winterocereus —> Cleistocactus Wittia KSch non Pantocsek > Pseudorhipsalis, Wittiocactus > Pseudorhipsalis, Yavin eee. e 2808 327 ‘Yungasocereus ++ .280 @ 197 Zehninerella > Facheiroa Zygocactus > Schlumbergera8 Artificial Keys to Genera Artificial Keys to Subfamilies and Genera For abbreviations in bold type see page 13 Key to Subfamilies Bo virally leafless (apart from floral scales), though the stems sometimes flattened, leaf-like; sd black or brown, sally exalt ay whol or ary enveloped by arming she or itu with cork srphiok or eopti- ‘lar pad; glochids absent o = Bo with functional leaves, atleast on new growth subfam, Cactoideae 2 2. Sd black or brown, usually exarillate, rarely wholly or partly enveloped by a mucilage sheath or the hilum with a corky strophiole or strophiolar pad; g} absent 3 — Sd encased in a usually whitish or brownish, bony, trichomatous, alveolate or wing-like aril; ar tufted with gl (minute- ly barbed spines), usualy in addition to larger spines 3. Botree-like, shrubby or scandent; Iv broad and flat (trop. America) — Bo low caespitose shrubs; Iv terete (Andes of $ Argentina, $ Chile) subfam, Opuntioideae (key 10 genera on page 13) subfam. Pereskioideae (Pereskia) subfam. Maihuenioideae (Maihuenia) Key to Subfamily Cactoideae 1 Br flat (leaf-tke) or 2-winged, atleast in part; or if 3-S-ribbed ‘or cylindric then <12 mm @ and not continuously ribbed; usually spineless or with fine bristles only ...... Group A — Br 3.4-winged or ribbed, or else S-many-ribbed or tubercled 2 2. Br scrambling, trailing or pendent, slender, often segmented, often emitting’ aerial roots, 3-4-winged or angled, or 5-12- ribbed; sp usually weak and bristly or very short, or hone Group B — _Brerect or ascending, not scrambling, slender to very stout, not producing aerial roots, 3-4-winged or angled, or few- to many- ribbed, or tubercled; sp often well-developed 3 3. Br elongate, cylindric or columnar, 3-4-winged or angled, or ribbed, >2x as long as thick before flowering; f1 nocturnal or diurnal 4 ~ Br globular or depressed, ribbed oF tubercled, <2x as long as thick (above ground) when flowering; 1 mostly diurnal .....7 Figar similar to the non-flowering, of less spiny 3 = Flgar different from the non-flowering, more hairy and/or bristly Group F 5. Pe with spines, bristles, hairs or felt; se conspicuous to obsolete 6 = Pe (at anthesis) nearly naked, or with se only Group E 6, Pe with spines or bristles Group C = Pe with hair-spines or hairs Group D 7._ Br ribbed, or ifubercled the tubercles disposed in more or less vertical rows 8 ~ Brovbercled the bers spill isponed, or er rarely both ribs and tubercles absent Fl arising in terminal cephalium “Grow ¢ enh not developed (but the brancirapex sometimes densely woolly) 6 Pesan bp beating cols wih wo, bis spines Group H = Peand hyp naked or with hairless se only Group J 10. Tub somewhat leaflike or scalelike, arranged as if in a rosette or cone Group K Tub not leaflike, usually gibbous, or absent Group L Group A [Rhipsalideae, Hylocereeae and Echinocereeae in part) 1. Flusually <# em @; hyp very short or none, rarely <5 em, or br divided into short segments <8 em 2 = FI large, usually >10 cm long: hyp well-developed; br not divided into short segments 10 2, Fl-<2em long, creamy white, arising atthe edges of the brseg, 3 ~ 52m ong, tightly colored (atest te sje) aisng ator near the tips ofthe brseg. 9 3. Br slender-cylindric or angled 4 = Br flat (2-winged), atleast in part 6 4 Brseg mainly arising singly from the sides of older ones, never in apical clusters 3 Brseg mainly arising at the apices of older ones, often in clus- ters, or main br producing short lateral segments only 7-15 mm. (R mesembryanthemoides) -Rhipsalis Pe with minute scales and areoles; fr when mature usually ‘translucent, veiny: frar with bristly spines « Pheiffera Pe usually naked: fr opaque, naked Lepisminm ‘Adult br all subsimilar, or if of 2 forms then the flat segments coarsely toothed (L. houlletianum); $d <1.5 mm, black or brown 6 ‘Adult br of 2 forms, the main axes terete atleast basally, the lat- eras fat, crenate; sd usually 1.5-2 mm, black Pseudorhipsalis Brseg mainly arising singly from the sides of older ones, never in apical clusters 8 Brseg mainly arising atthe apices of older ones, sometimes 2 or ‘more together | Rhipsalis Pe with minute scales and areoles; fr when mature usually +translucent, veiny, rem with large abscission zone .. Pfeiffera Pe usually naked; fr opaque; rem with small abscission zone Lepismium Fl regular; hyp <5 mm; lowermost sta not forming a ring round the style-base Hatiora Fl zygomorphic; hyp 8 mm or more; lowermost sta forming a ring round the style-base Schlumbergera FI diurnal, pink or red Disocactus Fl nocturnal, whi 10 Ar of flat br spineless 2 Ar of flat br spiny B Peshyp spineless Epiphyllum Pe+hyp with bristly spines + Selenicereus Bo epiphytic or lithophytic: br clinging by adventitious roots ‘Strophocactus Bo terrestrial shrubs; br not producing adventitious roots ‘Acanthocereus Group B [Hylocereeae in part; Echinocereeae in part] 1 2, Fi nocturnal, white, pale yellow or pale pink Fl diurnal, yellow, orange, red, pink or purple Bo creeping or climbing, often emitting aerial roots Bo shrubby, prostrate or scrambling, not rooting aerially (except ‘occasionally at tip) but rootstock often tuberous 6 Pe with conspicuous triangular se, enlarging in fruit; br usually 3-winged; sp very short Hylocereus Pe without conspicuous triangular se; br and sp various ....4 Fl shortly funnelform; hyp <5 em Weberocereus Fl clongate-funnelform; hyp >5 cm 3 Ri 3-10 wing-like, 2-15 em broad ‘Strophocactus Ri (2)5-12, low or very low SelenicereusArtificial Keys to Genera 9 R napiform or tuberous and Dahlialike; br slender, often minutely pubescent, dark-coloured and striate or speckled, not rooting aerally; sp generally very short, appressed Peniocereus R not tuberous, of tubers not as above; br various, not striate or speckled, sometimes rooting aerially; sp well-developed or short and weak 7 Br sharply 3-S-winged or angled; hyp stout, rigid, markedly fared; se scattered, felted and sometimes spiny but not hairy in their axils ‘Acanthocereus Br 3-4-angled or with 5-12 low ribs; hyp relatively slender, not rigid or markedly flared; se numerous, hairy in their axils Harrisia Sti green . Echinocereus Sti not green .... ee 9 Finearly rotate; hyp and tep very short Lapoceren (sala Coronas sp) Fl funnelform: hyp and tep elongate ... i 10 Br 3-5-winged, angled or ribbed .... * Disocactus Br very slender, with 6-12 low ribs u R tuberous, turip- or Dahlia-like; l white or epidermis minute- ly papillatehairy as Peniocereus R not tuberous; fl coloured: epidermis smooth or pitted... 12 ‘Ar <1.4-em apart on the ribs... B ‘Ar 1.4-2.2 em apart on the ribs . .. Echinocereus (E. pensilis) Brseg elongate, >30 cm; f= 2ygomorphic Disocactus Brg shor, wally <10 co: reget Echinopsis (E. chamaecereus) Group C [Cereoids with spiny or bristly pericarpel; no cephalium) 1 Per concealed during bud-development; br 3-4-ribbed, light, green Calymmanthium Per visible during bud-development; br various 2 Figar becoming elongate, proliferous, densely felted; i 4-8 -. Neoraimondia Fgar remaining cushion-like, not proliferous: Fi 3-30 or more 3 Br segmented or with abrupt constrictions; per short 4 Br not segmented or constricted: perianth-limb various ..... 5 FI small, -4 em (Cuba, Hispaniola, Puerto Rico) Leptocereus Fl larger, 6-11 m (Colombia, Ecuador, Peru) .. Armatocereus Bo low shrubs <1-2 m or unbranched: br <6.em@ ........6 Larger shrubs, or tee-like or columnar plants 2 Offsets and buds bursting through the epidermis; sti green Echinocereus Offsets and buds not bursting through the riders tl ot bright green ceed Fl narrowly tubulay-funnelform 8 Fl rotate-campanulate 10 Fl orange-red (Mexico: Baja California Sur) Echinocereus (E. pensilis) 9 Fl white Bo erect, <2(-3) m, rarely branched (E Brazil) ... Leocereus Bo low, shrubby (Galipagos Islands) Brachycereus Ri 10-25; fl yellow of white un Ri 5-10; fred or yellow Corryocactus Ri 20-25 (Mexico: Baja California; adjacent USA) css Bergerocactus Ri 10-13 Pera) Mila Fl campanulate, diumal (Peru, Bolivia, Chile) 13 Fl various, not campanulate, mostly nocturnal 4 ‘Ar conspicuously woolly or hairy; fl white or pale pink Eulychnia ‘Ar not woolly or hairy: brightly coloured .... Corryocactus Finarrowly tubular, bright red, 4-12.em, with oblique limb (NW Mexico) ‘Stenocereus (Rathbunia) Fl various, not bright red, regular 15 15, Hyp longer than pe 16 =" Hyp very short Polaskia 16, Fltubular with narrow limb; ri 4-6; sta throat-circle present ‘Samaipaticereus — Fl various; ri (5-)6-30 or more; sta throat-circle absent... 17 17. Figar confluent or discrete; tubular, with narrow perianth: transition from pe to hyp indistinct; ar of pe and hyp with bris- tes and hairs Pachycereus = Figar discrete; 1 tubular-funnelform, funnelform or campanu- te; anton om peo hyp sully abr: of hyp except the lowest, without bristles or hairs 18, Freubularfonnelform: be columnar fewbrancheds Hi? o more ‘Neobuxbaumia (N. te. — FI funnelform or campanuiate: bo shrubby or tree-like; ri 4-13-19 in S. hurber’) Stenocereus Group D [Cereoid Trichocereeae, with Eulychnia in par; no cephal- ium} 1. Hyp very short; ri 9-17 Eulychnia = Hyp well-developed to elongate 2 2. Sta inserted in a single series. Haageocereus Sta inserted in two series 3 3. Per narrow; fl nocturnal or diurnal, white or coloured 4 = Per broad; fl nocturnal, white 6 4. Fl nocturnal, white « Facheiroa = Fi diumal, mostly red (humming-bird syndrome) $ 5. Ar not clothed with long white hairs; fr 1~4 cm @, pulpy. inde- hiscent or bursting at one side Cleistocactus = Ar usually clothed with long white hairs (hair-spines): fr hollow; sd escaping at base, or if pulpy then 4-5 cm Oreocereus 6. Br often stout, not segmented; habit various (S America) <<. Echinopsis (see also Harrisia) = Br slender-elongate, 1-2'm x 2-5 em, or shortly segmented (SE & Cent. W Brazil) vce Arthrocereus Group E [Cereoids with scaly or naked pericarpel; no cephalium) FL very small, <2.5 cm, <9 per areole; ri 5-9 Fl Larger, 2.5 em or more, usually solitary; ri 3-30 or more .. 2 -Myrillocactus I 2. Peand hyp more or less scaly ceed 5, Reand yp naked or netrly so 10 3. Se chanaceous beans Escontria = Se not chartaceous 4 4 Scot hyp distant, butte bases sometimes strongly decuret 5 = Sc of hyp imbricate 5. Senot decent unelonm or salveronn, 418 om 26 = Se strongly decurrent; fl tubular to funnelform-campanulate, 25-8em 7 6. Fisalverform, 49 cm (Galapagos Islands). Jasminocereus = Fifunnelform, 12-15 em (Bolivia, Argentina) Stetsonia 7. Fr pulp red, relatively juicy Carnegiea = Fr pulp white, not juicy Neobuxbaumia 8 labor tobular-carpannate; st throa-crcle present (Bra) : Brasilicereus Fl ubular or wubular-funnelform: sta throat-circle absent (Peru, Bolivia, N Chile) es) 9, Fi diurnal, pink; flar white-felted Castelanosia Fi nocturnal, whitish or greenish; flar naked ..... Browningia 10. Fr red or yellow, often slightly blue-waxy; br-tissue not darken- ing when cut . u = Fr intensely blue-waxy; br tissue darkenshg when cut ceeeeee Cipocereus 1, Flapical Pierrebraunia =) Fllateral Cereus10 Artificial Keys to Genera Group F [Cereoids with modified flowering zone or cephalium, and ‘Neoraimondial |. Flgar elongating when old, flowering repeatedly ‘Neoraimondia = Figar not elongating, flowering once only 2 2. Ceph discontinuous, altemating with vegetative growth and marked by rings of bristles 3 ~ Ceph apical or lateral, continuous or discontinuous but not ring like wo 3. Br slender, constricted at each flowering zone 4 = Br columnar, not constricted at the flowering zones « Cephalocereus (Neodawsonia) (see also Haageocereus acranthus) 4. Fl>3.em @, white: hyp green; fruits blue-green; ri 13-18 ‘Stephanocereus (S. leucostele) = FI cm @ red, pink or yellow; fruits red, pink, purple or brownish; ri 7-14 ‘Arrajadoa 5. Ceph covering the whole apex 6 Ceph restricted to one side or discontinuous «2.2.0.2... 10 6. Br bottle-shaped, <1.5 m, usually unbranched; white within ‘Stephanocereus (S. uetzelburgii) = Br large shrubs or tree-like plants, not bottle-shaped: if less than 40 cm, Med 7 Ceph composed of long or very dense briste-spines 8 Ceph composed of wool and hairs or bristles Oreocereus (O. doelzianus) (see also Leptocereus paniculatus) 8. Pe with imbricate scales Browningia Pe with decurrent scales cose 9, RESIS Pachycereus = Ri17-26 ‘Neobuxbaumia (N. macrocephala) 10. Flowering zone not clearly defined, but the fl immersed in long woolly hairs Pilosocereus ~ Flowering zone or eeph clearly defined by massive development ‘of matted wool and/or bristles. ll 11, Peand fr scaly, the se conspicuous or minute, hairy i their axils 2 = Peand fr naked, or if with minute se these haitless in their axils 15 12, Fl diumal, brightly coloured Cleistocactus spp. (see also Denmoza) Fl nocturnal, pale or dull coloured 1B 13. FI produced only on stems ¢. 5 m; pe and hyp with minute, almost hairless se Cephalocereus = Fl produced on stems 2 m or less: pe and hyp with conspicuous se or hairs 4 14, Br masked by dense white spines and/or hairs; Br-tissue not dis- colouring on exposure to air, fl tubular-campanulate; se not imbricate Espostoa = Br green, not masked by dense spines and/or hairs; br-tissue turning brownish when exposed to air; fl tubular; se imbricate Facheiroa 15, Fr depressed-globose to broadly ovoid, depressed atthe point of attachment of the dried perianth, or laterally dehiscent, various- ly coloured 16 = Frobovoid-clavate to elliptic or globose, not depressed at apex. red, sometimes basally dehiscent, or < 15 mm O oS %6, lowering zone wih acospicuus hairs, but not foming a dense mat a ~ Flowering zone a dense mat or fleece-like ceph on one side of the br only Espostoopsis 17. Br highly mucilaginous (Central & § America) .. Pilosocereus ~ Br lacking mucilage (NW Venezuela) Cereus (C. mortensenii) 18, Fr not expressed from the ceph, indchiscent; rem tardily decid uous... Micranthocereus = Fr expressed from the ceph, opening by a basal pore; rem per- sistent Coleocephalocereus Group G [Globular genera with terminal cephalium) 1 Fl diumal, not fragrant, usually pink or red and less than 4 x 2.5 ‘em; fr indehiscent ‘Melocactus = Fl nocturnal, fragrant, white, 4-9 x 4-8 cm; fr dehiscent by lateral splits Discocactus Group H [Notocacteae, Trichocereeae, Cacteae in part; Echino- ‘cereus spp.) 1. Pe and hyp bristly or spiny; lb erumpent and/or sti green and fr juicy to Neshy Echinocereus — Pe and hyp not spiny, or the hyp with some bristles; lb not cerumpent; sti not green, or if green then fr not juicy or fleshy 2 Fi regul 3 Fl regular or bilaterally symmetric, lacking a sta throat-circle cesevsvetsesteeess 5 Fi mostly lateral, <4 em, variously coloured Echinopsis Flapical to subapical, 2.5-4(-6) cm, yellow to reddish orange 4. Br globose or somewhat elongate, simple or sparsely branched, not densely spiny Rebutia — Br cylindric, freely branched, covered with bristly spines . Mila Flat the shoulder of the br or below Rebutia = Flnear the br-apex 6 6. Se of hyp (atleast the uppermost) with wool and bristles in the axils cee 7 = Scot hyp only hairy or woolly in the axils a 7. Fr woolly; apical se spinescent Eriosyce Fr only sparsely woolly, or glabrous; apical se not spinescent 8. Frclavatew oblong, expanding and becoming partly hollow at maturity, opening atthe base; sd with relatively small hilum and no strophiole; testa often folded and ridged... . Neoporteria — Fr globose to ovoid or cylindric, indehiscen,spiving vertically ‘or disintegrating regularly: or if clavate and expanding then the 4 witha broad ium (equaling the diameter of he ee) and strophiole 9 Brdark reddish brows wt surace round and sometines ‘obscured by waxy whitish se; large mocilage bodies present in the cortex or with conspicuous mucilage ducts (Brazil: Minas Gerais) Uebelmannia = Br not as above: epidermis green or br <4 cm @; fr not usually red 10 10. Sp never hooked: f either cleistogamous or more than half the diameter of the br; sd with depressed hilum and no strophiole Frailea = Sp hooked or not; fl never cleistogamous, less than half the diameter of the br: sd not as above Parodia 11. Sc of pe and hyp, and apex of outermost tepals, narrow, sharply pointed, light o dark brown: fl various, regular, not tubular nor scarlet 12 — Se and outermost tepals not as above; fl various, sometimes bilaterally symmetric and/or scarlet 15 12, Bo more or less spotted or covered with small tufts of whitish ti- chomes; sd cap-shaped, with deeply sunken hilum; f1 yellow, sometimes red in the throat ‘Astrophytum — Bo and sd not as above; various : 13 13, Lower esp recurved at tip, brownish, very stout, transversely ridged: fr bright red Echinocactus (E. texensis) Cp nearly straight, or not as above; fr not red «....-..... 14 14, Fl yellow or pink; pe densely woolly, with se atthe apex only Echinocactus Fl pink or white; pe not densely woolly, scaly throughout Echinopsis 15. Fl tubular, per not expanded; sta and sty protruding: fr spliting laterally when ripe, Neshy inside DenmozaArtificial Keys to Genera 1 = Fl not as above: fr opening at or splitting from the base, dry inside 16 16, Flregular; fr opening at base Oroya = Flusually bilaterally symmetric; fr splitting laterally or from the base Matucana Group J [Notocacteae, Trichocereeae and Cacteae in part] |. Br ridged and wrinkled between the ri; sd minute, 0.0.7 mm, strophiolate Aztekium = Brand sd not as above 2 2. Bo spineless; ar tufted with woolly hairs only. - Lophophora = Bo sspiny 3 3. Flregular 4 = Fibilaterally symmetric: hyp curved ‘Matucana 4. rey. opening at apex, the base ofthe dried perianth detaching like a lid: 1 yellow: hyp very short Copiapoa ~ Fr dry or juicy, opening basally or laterally or disimegrating imegularly or indchiscent 5 5. Ri very thin, more or less wavy, or if few then upper sp 1-3 large, flattened, lower 1-2 much smaller terete ... Stenacactus = Rinot thin and wavy: sp not as above 6 6. Fr fleshy, usually bursting laterally; ri usually ‘chinned’ beneath the arcoles; areolar glands absent Gymnocalycium = Fredry, or juicy but indehiscent; i not “chinned'; areolar glands often present 1 7. Mature br large, more than 20 em @, or smaller and the sd pitted; ri well-defined Ferocactus — Mature br 4-20 em @; sd smooth, reticulate or tuberculate; ri poorly defined, strongly tubercled 8 8. Sp obscuring the br. or the esp strongly recurved to hooked: fl never yellow with red throat; sti sometimes green; hmr lateral or, oblique to main axis of seed Sclerocactus ~ Sp not as above, or ft yellow with red throat; sti never green; ‘hme terminating long-axis of seed Thelocactus Group K [Cacteae in part; stems with leaf- or scale-ike tubercles} |, Tub very long (to 15 em) and thin, 3-angled; sp long, papery Leuchtenbergia = Tub various, ot as above; sp 0 or caducous or bristle-like .. 2 2 Tu rater iin, angular, seale-ike, closely overaping: conspicuous = Tube, tiguetros and sult or spyeamial, aot closely ‘overlapping: sp inconspicuous or absent Ariocarpus 3. Fl purplish pink; sta not sensitive; tub appressed throughout, as ina bulb Pelecyphora (P.strobiliformis) = Fipale yellow; sta sensitive: tub, except the youngest, spread ing Obregonia Group L (Notocactese, Trichocereeae and Cacteae in part] 1. Brbutton-like,c. 1 em @, without rior tub Blossfeldia = Br tuberculate 2 2, Ar situated between the tubercles; tubercle apex naked ‘Neowerdermannia — Ar situated at the tubercle-apex, sometimes extending via a ‘groove 10 the ‘axil, or bi-panite (with a discrete axillary Portion) 3 3. Flarisng a the tips of the tubercles. or in short groove towards, the ‘axil” 4 Fl arising in the tubercular ‘axils" 19 Fl bome at or near br-apex 3 Fl borne on the ‘shoulder’ of the br or below Rebutia Pe and fr bearing se, wool and sometimes bristles: fr dry inside, indehiscent, disintegrating irregularly or partly hollow 6 ~ Beand fr nuked or besrng hes nls juey,ndeisent, ‘or dry. opening at base, apex or laterally 8 2B, 24 FI yellow, but often cleistogamous, the perianth then scarcely developing: sd 1-3 mm, shiny, with sunken hilum .... Frailea FI yellow, red, pink or whitish, never cleistogamous; sd, 0.5-1.2 mm, shiny or dul, the hilum not sunken 7 Fr more or less globose, 5-10 mm; sd 0.5-1 mm, strophiolate Parodia Fr clavate, 10-25 mm; sd 0.8-1.9 mm, not strophiolate Eriosyce Fr opening near apex 9 Fr indehiscent or opening laterally or below, the dried perianth- base or its scar remaining firmly attached to the pericarp on all sides cell Fr sunken and hidden in the depressed apex until the next fg season; rem deciduous (Argentina) Yavia Fr not hidden; rem persistent, atleast until dehiscence... 10 Fr splitting laterally; r fibrous: fl pink, white or yellow; sd 1.5-5 mm (USA) Pediocactus Fr cup-like, sometimes splitting longitudinally also; r tuberous: £1 yellow, rarely reddish; sd 0.8-2.1 mm (Chile) . ... Copiapoa Pe and fr always visible, the latter dry, opening laterally or basally, dll green or brownish 2 Pe hidden between the tubercles or by spines and wool at flow- cering-time: or fr juicy. indehiscent, bright green, pink or red, oF dry and remaining hidden when ripe 15 ‘Sp flatened, resembling dried grass leaves; bo clavate-cylin- dric; sti green Sclerocactus ‘Sp not as above or br depressed-globose; sti not green... 13, Pe and fr naked, rarely with 1 or 2 se; fl to 3.5 x 4 em; sd 1-2 mm, black, not strophiolate Neolloydia Pe aod fr wih 2 of more se: <6 8 cm ors only 0.5m, brown, strophiolate ...... 4 Fr opening by a basal pore: sd 1.5-2.5 mm, black, not strophio- late; p strong or dense Thelocactus Fr opening by 1-3 vertical spits; sd 0.5 mm, brown, strophio- late: sp few, weak, caducous Strombocactus Bo spineless; ar tufted with woolly hairs only ... Lophophora Bo spiny = 16 ‘Tub conspicuous, >1 mm; areolar groove short or extending 10 the base "7 ‘Tub minute, ¢.1'mm, not grooved Epithelantha Fr juicy, often brightly coloured; sd black or brown; testa smooth or pitted, 18 Fr dry, dull-coloured: sd black; testa tuberculate . . Neolloydia ‘Testa cells tabular; otep entire, or areoles with glands Coryphantha Testa cells tabular-concave or par-concave (pitted); otep fringed; areolar glands absent ‘Acharagma ‘Tub low, pale grey-green, punctate; 2-3 em, yellow, with deep green sth ‘Ortegocactus Not as above 2 20 Figar grooved or ridged between the sterile apical and fertile axillary portions <2... eee eceseseeseeterestseee 21 Figar bi-partte, the apical and axillary portions discrete... 24 ‘Tub hatchet-shaped; sp pectinate Pelecyphora (P.aselliformis) Tub cylindric, conic or gibbous; sp various 2 3. Testa cells tabular; otep entire, or ar with glands 23 Testa cells tabular-concave or par-concave (pitted); otep fringed; areolar glands absent . Csp 3-4 hooked; fr slender-eylindric, scented .... Cumarinia sp straight or curved, rarely 1 hooked: fr globose t0 obovoid, not scented Coryphantha Sd black; celis of testa tabular or somewhat convex ‘Mammilloydia ‘Sd black or brown: if black the testa cells par-concave (pitted) ‘Mammillaria12. Artificial Keys to Genera Key to subfamily Opuntioideae Based on the key by Wolfgang Stuppy (Succ. Pl. Research 6: 41-43. 2002) Br with persistent broad, flat leaves 2 Br with small terete, mostly early caducous leaves 3 Br segmented; fl red or pink, usually terminal; s@ 410 mm (South America) ‘Quiabentia Br not segmented; fl yellow, usually lateral; sd 4-5 mm (Mexico, Guatemala) Pereskiopsis Br globose or cylindric 4 Br compressed (at least the terminal ones) 15 Sp with a (sometimes rudimentary) papery sheath (North America) . Sp without papery sheath (South America). ; Bo small trees and shrubs; br with indefinite growth and prom nent longitudinally elongated tubercles; sp mostly rounded with the epidermis separating completely in a decidous papery sheath gl flattened at base Cylindropuntia Bo shrubs or low, cushion-forming plants; shoots with determi nate or indie growth p flatened with ony a rudimentary papery sheath; shape of glochid base rounded Bo shrubby, thicket forming: brseg terete, 8-10-ribbed with indefinite growth; flgar without glochids Grusonia Bo low shruts, articulated, cushion-forming: brseg clavate, not ribbed; flgar with glochids « Corynopuntia Br not segmented, growth indefinite 8 Bo segmented; brseg with determinate growth 9 Bo shrubs or small ees; lv often >4 mm, caducous - ‘Austrocylindropuntia Bo clambering or scrambling, sparsely branched shrubs; IV ‘minute, early caducous (E Brazil) Tacinga funalis Bo low shrubs, often creeping and rooting, segmented; brseg small, determinate, globose or cylindric to compressed or sub- terete: fr juicy, but with only litle pulp, dehiscing by a single lateral split, the two halves then open wide apart; sd 2.5~4 mm, irregular reniform, outer part of funicular envelope intensely red and juicy when fresh, rugosely sculptured and brown to black when dry Tunilla Bo cylindric, never compressed: fr and sd not as above... 10 Bo small geophytes with tuberous roots; brseg globose or cylin- dric; glochids with distinctive basal’ structure (see generic ‘description; apical; fr dry, capsular, transversely dehiscent in the lower third: sd strongly compressed, withthe funicular girdle Cari”) developed into a papery wing (Argentina) Fl not apical fr not capsular: sd not winged " ‘Ar deeply sunken into globular to pyriform cavities with small, ‘openings: fr dry, irregularly dehiscent, without pulp; sd with large, spongy funicular aril; bo fragile shrubs; br often monil form, brseg globose, cylindric or obovate; (Argentina) Tephrocactus Ar not sunken into globular to pyriform cavities: funicular ail 12, 13, not spongy 2 Bo a squarrose shrub, 0.5-1.5 m: br cylindric, segmented; brseg 7-20 x 3-6 em, conspicuously tuberculate, strongly glaucous (especially when young) (Chile) ‘Miqueliopuntia Bo low growing, either forming compact cushions or very small individuals composed of only 1-5 obconic or cylindric seg- ‘ments; funicular girdle not triangular in cross section 15 Bo small; brseg 1-5 obconic or cylindric; sp pectinate; fr dry: frar without glochids ‘Maihueniopsis Bo low-growing, forming compact, semiglobose cushions; br ‘segmented; brseg globose to ovoid, ellipsoid, obconic or cyl dric; sp not pectinat; fr not dry; frar with glochids 4 Brseg with ar evenly distributed; fr juicy, with pulp: sd white to yellow, laterally flattened, lenticular to irregularly angular, woolly ‘Maihueniopsis Brseg with ar crowded towards the apex; fr thick-walled, with the sd lying dry in the pulp-free loculus: sd globular to pyriform, ¢labrous or with a few inconspicuous trichomes; embryo hook- shaped; perisperm reduced ‘Cumutopuntia Bo arborescent, developing a distinct trunk 16 Bo small to large shrubs (but some Opuntia spp. arborescent) : ve 7 ‘Ultimate brseg large, compressed, curved, distinctly inequilater- al 1 with pe elongate, sometimes curved like the terminal brseg; style; per small, usually orange or red; sty with a basal thicken- ing: sd small, woolly (Florida, Caribbean) Consolea Ultimate brseg thin, lea-like but spiny, deciduous; with aring of hair-like staminodes between the tepals and the stamens; sd very large, 6.5-10 mm @ (South America) .... Brasiliopuntia, Bo low shrubs, often creeping and rooting, segmented; brseg, small, determinate, compressed to sublerete; fr juicy, but with only little pulp, dehiscing by a single lateral split, the two halves then open wide apart; sd 2.5-4 mm, irregular reniform, outer part of funicular envelope intensely red and juicy when fresh, rugosely sculptured and brown to black when dry (South America) Tunilla Bo shrubs or trees; brseg compressed (very rarely cylindric), orbicular, obovate to elliptic; fr indehiscent, with watery-juicy pulp: sd not brightly coloured and juicy on the outside, surface never rugose 18 Fl almost completely closed during anthesis: per erect; sta and sty exserted: pg without reticulate ektexinous omamentation (Mexico to Panama) Nopalea Flopen during anthesis: per spreading to rotate 19 Pg with reticulate ektexinous omamentation; sd laterally com- pressed; embryo spirally enrolled; sta often sensitive, closing round the sty if touched Opuntia Pg without reticulate ektexinous omamentation; sd globose to pyriform: stamens not sensitive (E Brazil) - Tacinga13 (Conventions and. Abbreviations Format of catalogue entries ‘The entries mostly consist of two or three paragraphs, the first con- taining name, source and distribution data, the second the description (for accepted species and subspecies) and the third any relevant com ‘ments. The data in the fist and second are given in a standard order, with each main item preceded by an abbreviation in bold type. The commentary paragraphs are preceded by a ‘bullet’ (+). ‘Name and distribution data for genera: Genus name, abbreviat- cd authority and literature citation. Etymology. Basionym if applica- ble, with data as for accepted name. Type species. Synonyms, Distribution to country level (2-etter codes: see list) Literature. ‘Name data for species and subspecies: Record no. The unique S- Aigit record number of the entry in the lexicon database. Species name, abbreviated authority, year and place of publication (see expla- nation below). Basionym, if applicable, with its record number and reference. Type: country code, state, department or province, locality, altitude, other details, date of collection, collector(s). number. acronym of herbarium where deposited. Distribution (if known range extends beyond the country or state of origin ofthe type) Authority abbreviations (pages 14-16) [Names of sole authors are abbreviated (if necessary) to a maximum of five letters (no full-stop), joint authors to a maximum of 11, including a “+’ between them: ifthe citation includes more than two authors, the normal abbreviation ofthe first author only is given, fol- owed by "+". Accents and diacritical signs are not used. References (for book and journal abbreviations see pages 17-19) Scope. Normally only the first place of (valid) publication is given, ‘but supplementary references may be given if they are an important source of data. Content. Year of publication, followed by a journal or book acronym or abbreviation, volume andior issue number (if any). page number (or ‘[unpag]’) and plate or fig. number(s) or ‘illus’ to Indicated the article is illustrated; when par of a volume, the issue ‘number may be indicated in brackets °( )° immediately after the volume number. Punctuation. Year and place of publication are sep- arated by a slash (/"); volume/part numbers are followed by a colon (2) then the page number(s); if there is no volume number, book abbreviations are followed by the page number (no comma). Name and Type data Deseriptions (Descr.) Article 8 of the International Code of Botanical Nomenclature defines various cat- egories of "types. Abbreviations for these used in NCL are listed below along with other abbreviations and symbols used in NCL entries. For the acronyms of herbaria where types are deposited, readers are refered to Index Herbariorum. The name-data for non-recognized taxa (names in italic type) are normally fol- lowed by an arrow symbol °->" meaning “referred to" or ‘synonym of” (see note ‘on treatment of synonyms on p. 2). The symbol "= "is occasionaly used to indicate “equivalent to" or ‘the same as’, and (for provisionally recognized taxa (bold italic type) the symbol * ~ "to mean ‘perhaps referable 1". B_basionym Syn synonym(s) 1nd not designated Distr distribution T —typeltypification np_not preserved ET epitype TL typelocality nx. not extant Etym etymology Mus illustration(s)* holo. holotype = equivalent (name) ur iso. isotype © name best abandoned Lit lecto. lectotype @ outlawed name Nr neo. neotype > referred 10 st para. paratype x. hybrid * Only for references to important illutration(s) not in the NCL Atlas Distribution: Country Codes The two-letter codes mostly follow the International Standards Organization (1SO) list. Names of other countries are given in ful AN Netherlands Antilles GT Guatemala ‘TT Trinidad & Tobago AAR Argentina GY Guyana US United States BO Bolivia HN Honduras UY Uruguay BR Brazil HT Haiti VE Venezuela BS Bahamas JM Jamaica VG British Virgin Iss. BZ Belize KY Cayman Islands. VI_US Virgin Islands CA Canada MX Mexico WI Lesser Antilles CL Chile NI Nicaragua XC from cultivation CO Colombia PA Panama CR Costa Rica PE Peru dept. Departamento CU Cuba PR Puerto Rico mpio Municipio DO Dominican Republic PY Paraguay prov. Provincia EC Ecuador SR. Suriname GF French Guiana SV BI Salvador cult, cultivated Coordinates of latitude and longitude may be given thus: 28°S, or (when more precise) thus: 16:128/56:20W or 6:41:25.58/76:18:17.5W. In genera with subdivisions, descriptions of species are preceded by the number of the relevant subdivi- sion, in ‘[ |’ (bold type if this also relates to the arrangement of the illustrations in the Atlas volume, and the species i illustrated there). Abbreviations for the component parts of the plants are listed below, abrbbr branch types! ne nectar chamber anth anthers, tep outer tepals ar areoles pe percarpel ax axils Pe pescarpt bo bodyyhabit—per_—peranth br branches (stems) pg pollen-grains csp central spine(s) pulp fruit pulp fi filaments F root) ootstock A flowers) rem peranth remnant fla lowerareoles! rl ibs fh lowerbudis) rsp radial spines fg flowering se. scales fgar flowering areoes sd seeds) fe fruit Seg sezment(s) at lochids sp spines gsp/glsp areolar glands sta stamens fr glandspines sti stigma(s)obes hilum tilim Stipe basal sem hhmr hilum micropylar sty style region tep tepals) hyp hypanthium? testa testa, seedoat itep ier tepals tr trunk Wve leaves tub tubercles ms mucilage sheath 1. Used where branches are dimorphic (of two types): 2. Abbreviations may be compounded, as here and in “rseg"(branch-segments), ‘pes (pericarpel-scals) ete 3. The part ofthe receptacle tube above the pericarpel: 4. Here used fr the rind of the fruit Measurements Unless otherwise stated, measurements are of length, or length x breadth or diameter (B). > more than © upto oF more than << upto, less than more or less © diameter X by, as in long x broad