Illustrated Handbook of Succulent Plants

Series Editors: U. Eggli, H. E. K. Hartmann

Springer-Verlag Berlin Heidelberg GmbH Illustrated Handbook of Succulent Plants (Eggli/Hartmann Eds.)

Illustrated Handbook of Succulent Plants: Monocotyledons Ed. by Urs Eggli (2001)

Illustrated Handbook of Succulent Plants: Dicotyledons Ed. by Urs Eggli (2002)

Illustrated Handbook of Succulent Plants: Aizoaceae A-E Ed. by H. E. K. Hartmann (2001)

Illustrated Handbook of Succulent Plants: Aizoaceae F-Z Ed. by H. E. K. Hartmann (2001)

Illustrated Handbook of Succulent Plants: Asclepiadaceae Ed. by F. Albers and U. Meve (2002)

Illustrated Handbook of Succulent Plants: Crassulaceae Ed. by Urs Eggli (2003) Heidrun E. K. Hartmann (Ed.)

Illustrated Handbook of Succulent Plants: Aizoaceae A-E

With 54 Figures and 384 Colour Photos, Printed in 48 Colour Plates

Springer DR. HEIDRUN E. K. H ARTMANN Institut fur Allgemeine Botanik OhnhorststraBe 18 22609 Hamburg Germany e-mail: [email protected]

With contributions by: C. 8ruckmann, M. Gerbaulet, S. A. Hammer, B. Hansen, H. E. K. Hartmann, H.-D. Ihlenfeldt, C. Klak, S. M. Pierce, M. Struck-Gerbaulet

Library of Congress CataJoging-in-Publieation Data

Illustraled handbook of sueeulent plants. Aizoaeeae / Heidrun E.K. Harlmann (ed.). p.em. Indudes bibliographical references. Contents: III A-E -- (2] F-Z. ISBN 978-3-642-62584-8 ISBN 978-3-642-56306-5 (eBook) DOI 10.1007/ISBN 978-3-642-56306-5 1. Aizoaceae--Classification. 1. Hartrnann, Heidrun E. K.

QK495.A32 144 2001 583'.53--d C21 2001040053

This work is subject to copyright. AII rights are reserved, whether the whole ar part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasling, reproduction on microfilms or in any olher way. and Slorage in dala banks. Duplication of this publicat ion or parts thereof is permitted only under the provisions of the German Copyright Law of Septernber 9, 196~, in its current version, and permission for use musi always be obtained from Springer-Verlag. Violations are liable (or prosecution under the German Copyright Law.

C Springer-Verlag Berlin Heide1berg 2002 Originally publishcd by Springcr-Verlag Berlin Hciddbcrg New York in 2002 Softcover rcprint of !he haroco"er ISI cdilion 2002 The use of general descriptive names.registered names, trademarks,etc. in this publication does nOI irnply, even in the absence of a specific statement, that such names are exempt from Ihe relevant prOlective laws and regulations and therefore free for general use.

Cover design: design & productioll GmbH. 69121 Heidelberg, Germany Typesetting: Data conversion by Buro Stasch, Bayreuth, Germany ([email protected]) 31/3111 - 5 4 3 2 1 - Printed on acid-free paper Preface

Members of the iceplant family, Aizoaceae, are appreciated by many people and disliked by some. Farmers value the plants for their browsing stock, hobbyists like to grow and flower mainly the "stone plants" of the family, and tourists enjoy the mass flowering of millions of annuals after good rains in the south western part of South Africa. Only taxonomists show some reservation against members of the Aizoaceae because of the size of the family and the confused and entangled systematics of many groups. For all groups of users, this book will offer some improvements. A worldwide web has been knitted among hundreds of interested people resulting in the pro• duction of this lexicon. As a consequence of the expanded collaboration over five continents, innumerable new results can be presented here, including many new delimitations of genera as well as new combinations and names of species. All conclusions are based on thousands of days spent on field work, thousands of original literature references inspected, thousands of herbarium sheets studied, thousands of scanning electron microscopic pictures compared, thousands of photographs taken, and thousands of taxonomic decisions made. As a result of all united efforts, an up-to-date survey of the state of the art can be presented, forming the basis for further research in the family and providing a challenge to every reader to add his special knowledge towards an improved second edition. This book continues in part the tradition of Jacobsen's "Handbook of Succu• lent Plants" vol. 3 (1960), and of the "Lexicon of Succulent Plants", published from 1955 onwards in different editions in German and English, in which Jacobsen treated up to 122 genera of Mesembryanthema under the heading "Mesembryan• themaceae" . In these two volumes dedicated to the family Aizoaceae, the Mesembryanthema are included. 234 genera have been named in the family Aizoaceae in the present circumscription, all of which are described here, but only 136 are accepted as valid. 29 new combinations, one new name, and two new species are published here; 189 taxa are lectotypified, 26 are neotypified. After a long trip in troubled waters, the manuscript was revised and updated in the year 2000, the closing date for inclusion of known publications being 31.12.2000. I am most grateful to all authors and colleagues who have helped to keep up with the numerous new descriptions, including contributions of taxa in the process of description now, and unpublished information. I should like to thank all collaborators and contributors of text or artwork. My special thanks go to Dr. Czeschlik and Dr. Lindenborn, Springer-Verlag, who made the publication possible at last.

H. E. K. Hartmann

Hamburg, September 2001 Acknowledgements

Maike Gerbaulet gratefully acknowledges travel grants from the Cactus and Suc• culent Society of America and the Mesem Study Group, England, constant sup• port by the staff members of the Bolus Herbarium, professional maintenance of her living plant collection by Ernst van Jaarsveld, and special support by Chikako Ueda. H. E. K. Hartmann is most grateful to about 500 officials, colleagues, friends, and students for all kindness and friendly support without which a work of this dimension would never have been possible. Over the 32 years of studies in Aizoaceae including field work in Argentina, Botswana, Ghana, India, Kenya, Lesotho, Mexico, Namibia, South Africa, Sudan, Swaziland, Tanzania, USA, and Zimbabwe numerous organizations and people have helped her work in many ways: giving permission to work in regions with restricted access like Diamond or other Mining Areas, Nature Reserves, Botani• cal Gardens, or Military grounds, getting permits from Nature Conservation Bod• ies for collecting protected plants in all countries visited, organizing sometimes extensive field trips, allowing to stay over night and feeding her well, searching for plants in sometimes rather inaccessable places and under difficult conditions, collecting data on distribution and taking her to places, keeping her company and helping during field work, driving cars and repairing them miraculously, finding very rare and difficult to trace literature, checking herbarium specimens, drying and shipping the collected herbarium material, doing scanning electron microscopic work, caring for living plants and growing them, measuring and draw• ing leaves, fruits, and flowers, turning drawings into ink versions for publication, discussing problems in finding roads or drawing the correct taxonomic conclu• sions, and, above all, being willing to share their knowledge with her. The directors of the herbaria B, BM, BOL, BOLO, HEL, HBG, K, NBG, PRE, UPS are thanked for most generous permission to study or get on loan valuable type material or photographs which made progress possible. B, BOL, and K are espe• cially acknowledged for very extensive loans over a much extended period. Financial support from the German Science Foundation (DFG) since 1969, from the Council of Scientific and Industrial Research in South Africa, the award of the Kistenbosch Jubilee Fund 1977, from the National Botanic Gardens via the Karoo Botanic Garden, the Universities of Cape Town and Pretoria, the Hansische UniversiHitsstiftung of the University of Hamburg, the University of Hamburg, and the Cactus and Succulent Society of America is most gratefully acknowl• edged. H.-D. Ihlenfeldt should like to thank the German Science Foundation for fi• nancial support over many years and the director and staff of BOL for the exten• sive loan of valuable herbarium material. 1. Niesler is grateful to S. Hammer for helpful information and material, to H. E. K. Hartmann for many discussions and support of her work. She should also like to thank the directors of BOL and K for extended loans of important material. N. Zimmermann gratefully acknowledges a travel grant from the Cactus and Succulent Society of America and the loan of material of Schwantesia from BOL. Contents

Introduction .... 1 The Family Aizoaceae ...... 1 Circumscription and Delimitation of the Family Aizoaceae ...... Subdivision of the Aizoaceae ...... 1 Diversity and Ecology of the Family ...... 2 Distribution ...... 2 Importance of Characters for the Delimitation of Taxa Within the Aizoaceae .. 2 Technical Information on the Handbook ...... 4 General Data ...... 4 Terminology for Descriptions ...... 4 Taxonomic Principles ...... 5 Rank and Limits of Genera ...... 5 Rank and Limits of Species ...... 5 Ranks Below Species ...... 6 General Scheme for the Treatment of Taxa ...... 6 Nomenclature and Typification ...... 6 Standard of Descriptions ...... 7 Orthography ...... 7 Keys ...... 7 Distribution ...... 7 Illustrations ..... 8

Aizoaceae ...... 9 Abryanthemum == Carpobrotus ...... 23 Acaulon == Aloinopsis ...... 23 Acrodon RUSCHIOIDEAE ...... 23 Acrosanthes AlZOOIDEAE ...... 25 Aethephyllum RUSCHIOIDEAE ...... 26 Agnirictus = Stomatium ...... 27 Aistocaulon == Aloinopsis ...... 27 Aizoanthemum AlZOOIDEAE ...... 28 Aizoon AlZOOIDEAE ...... 30 Aioinopsis RUSCHIOIDEAE ...... 33 Amoebophyllum = Phyllobolus ...... 37 Amphibolia RUSCHIOIDEAE ...... 37 Ancistrostigma = Trianthema ...... 39 Anisocalyx = Jacobsenia ...... 40 Anisostigma = Tetragonia ...... 40 Antegibbaeum RUSCHIOIDEAE ...... 40 Antimima RUSCHIOIDEAE ...... 41 Apatesia RUSCHIOIDEAE ...... 65 Aptenia MESEMBRYANTHEMOIDEAE ...... 66 Arenifera RUSCHIOIDEAE ...... 68 Argeta = Gibbaeum ...... 70 Argyroderma RUSCHIOIDEAE ...... 70 Aridaria MESEMBRYANTHEMOIDEAE ...... 75 Contents x

Aristanthus ::: Lampranthus ...... 80 Aspazoma MESEMBRYANTHEMOIDEAE ...... 80 Astridia RUSCHIOIDEAE ...... 81

Bergeranthus RUSCHIOIDEAE ...... 84 Berrisfordia ::: Conophytum ...... 86 Bijlia RUSCHIOIDEAE ...... 87 Bolusanthemum ::: Bijlia ...... 88 Braunsia RUSCHIOIDEAE ...... 88 Brownanthus MESEMBRYANTHEMOIDEAE ...... 91

Calamophyllum RUSCHIOIDEAE ...... 95 Callistigma ::: Mesembryanthemum ...... 96 Carpanthea RUSCHIOIDEAE ...... 96 Carpobrotus RUSCHIOIDEAE ...... 97 Carruanthus RUSCHIOIDEAE ...... 101 Caryotophora RUSCHIOIDEAE ...... 102 Caulipsolon MESEMBRYANTHEMOIDEAE ...... 103 Cephalophyllum RUSCHIOIDEAE ...... 103 Cerochlamys RUSCHIOIDEAE ...... 113 Chasmatophyllum RUSCHIOIDEAE ...... 115 Cheiridopsis RUSCHIOIDEAE ...... 117 Circandra RUSCHIOIDEAE ...... 129 Cleretum RUSCHIOIDEAE ...... 129 Conicosia RUSCHIOIDEAE ...... 131 Conophyllum ::: Mitrophyllum ...... 133 Conophytum RUSCHIOIDEAE ...... 134 Corpuscularia RUSCHIOIDEAE ...... 175 Crocanthus = Malephora ...... 178 Cryophytum = Mesembryanthemum ...... 178 Cylindrophyllum RUSCHIOIDEAE ...... 179 Cypselea SESUVIOIDEAE ...... 181

Dactylopsis = Phyllobolus .. 182 Deilanthe RUSCHIOIDEAE ...... 182 Delosperma RUSCHIOIDEAE ...... 184 Demidovia = Tetragonia ...... 210 Depacarpus = Meyerophytum ...... 210 Derenbergia = Conophytum ...... 211 Derenbergiella ::: Mesembryanthemum ...... 211 Dicrocaulon RUSCHIOIDEAE ...... 211 Didaste ::: Acrosanthes ...... 215 Didymaotus RUSCHIOIDEAE ...... 215 Dinteranthus RUSCHIOIDEAE ...... 215 Diplochonium ::: Sesuvium ...... 218 Diplosoma RUSCHIOIDEAE ...... 218 Disphyma RUSCHIOIDEAE ...... 220 Dorotheanthus RUSCHIOIDEAE ...... 222 Dracophilus RUSCHIOIDEAE ...... 225 Drosanthemopsis ::: Jacobsenia ...... 226 Drosanthemum RUSCHIOIDEAE ...... 227

Eberlanzia RUSCHIOIDEAE ...... 250 Ebracteola RUSCHIOIDEAE ...... 254 Echinus ::: Braunsia ...... 257 Ectotropis RUSCHIOIDEAE ...... 257 XI Contents

Enarganthe RUSCHIOIDEAE ...... 258 Erepsia RUSCHIOIDEAE ...... 258 Esterhuysenia RUSCHIOIDEAE ..... 267 Eurystigma = Mesembryanthemum .... 268

References .... 269

Colour Plates List of Abbreviations and Symbols

o ...... diameter nomen illeg. . nomen illegitimum, illegitimate ± ...... more or less name, not permitted under the > ...... bigger than ICBN ~ ...... bigger than or equal to nomen invalid. nomen invalidum, invalid name, < ... .. smaller than not fulfilling the requirements ::; ...... smaller than or equal to for valid publication as given by x ...... times; hybrid sign in a name ICBN ! ...... the cited herbarium sheet has been seen nov...... nova, novum, new = ...... equals; between names: heterotypic NT ...... neotype synonym, based on a different type p.a...... per annum, in a year than the accepted name of the taxon pers. comm ... personal communication == ...... homotypic synonym, based on the same p.p ...... pro parte, in part type as the accepted and valid name § ...... section S ...... seed(s) App...... appendix s.lat...... sensu latiore, in the wider sense Art...... article s.n...... sine numero, without a number ssp...... subspecies b ...... broad s.str...... sensu stricto, in the narrow sense c. . ... circa, about subg...... subgenus Chr ...... chromosomes t...... tabula, plate comb. nov..... combinatio nova, new combination t ...... thick Distr ...... Distribution T ...... holotype Ecol ...... Ecology var...... varietas, variety Etym ...... Etymology 2n ...... double chromosome set f ...... figure FI ...... flower( s) Abbreviations of names of authors of plant names Fr...... fruit(s) follows Brummitt R.K. & Powell C.E. (1992) Authors Gk ...... Greek of Plant Names. Royal Botanic Gardens. Kew in many cases, e.g. Haw. for Haworth, 1. for Linnaeus, or h ...... high N.E.Br. for N.E. Brown. Deviations have been found hort...... hortus, garden useful in cases in which the authors are rare within I ...... internode( s) the family. ICBN ...... International Code of Botanical Nomenclature Abbreviations of names of herbaria follows Holmgren in litt...... in litter is, in a letter et al. (1990) Index Herbariorum, Ed. 8 Inf ...... inflorescence( s) ISO ...... isotype(s) Names of genera are abbreviated by their first, capi• tal, letter within the text covering that genus. An ex• K ...... calyx ception has been made for M, in which case M. al• I ...... long ways stand for the genus Mesembryanthemum; L ...... leaf, leaves all other genera starting with an M are abbreviated La .... Latin by their first two letters, e.g. Ma. for Malephora or LT ...... lectotype Machairophyllum, respectively. List of Abbreviations and Symbols XIV

Abbreviations of Geographical Terms

Ab ...... Aberdeen Mo ...... Montagu A.B...... Alexander Bay MM ...... Mmabatho Ag .... Aggenys MP ...... Mpumalanga, Province Au ...... Aus Mu ...... Murraysburg B.E...... Barkly East N ...... north, northern Bi ...... Bitterfontein NC ...... Northern Cape Province BI ...... Bloemfontein No ... Noorduwer Br ...... Bredasdorp NP ...... Northern Province Bu . . . .. Burgersdorp NW ...... North Western Province B. W...... Beaufort West OR ...... Orange River Ca . . .. Carnarvon Ou ...... Oudtshoorn CI ...... Calvinia P.A...... Prince Albert Co ...... Carolina Pc ...... Potchefstroom Cr ... .. Cradock P.E...... Port Elizabeth C.T...... Cape Town Pi ...... Pietersburg E ...... east, eastern Pm ...... Posmasburg EC ...... Eastern Cape Province P.N ...... Port Nolloth E.L...... East London Po ...... Pofadder Pr ...... Pretoria Fr ...... Fraserburg FS ...... Free State, Province Rc ...... Richmond Ri ...... Riversdale Ga ...... Garies Ru ...... Rustenburg GAU ...... Gauteng, Province Gb ...... Gaborone S ...... south, southern Ge ...... George S Africa .. Republic of South Africa Gh ...... Grahamstown Sp ... Springbok Gr ...... Griquatown St ...... Steytlerville G.R ...... Graaff Reinet Su ...... Sutherland Sw ...... Swakopmund Ha ...... Harrismith Ho ...... Hondeklipbaai To ...... Touwsrivier Ja ...... Jagersfontein Ui ...... Uitenhage Jo...... Johannesburg Urn ...... Umtata Un ...... Uniondale Ke ...... Kenhardt Up ...... Upington Ki ...... Kimberley K.K...... Kap Kruis Va ...... Vanrhynsdorp KN ...... KwazuluNatal, Province Vr ...... Vryburg Ku ...... Kuruman VR ...... Vaal River La ...... Laingsburg W ...... west, western L.B...... Lamberts Bay Wa ...... Warmbad Lo ...... Loeriesfontein WC ...... Western Cape Province L. T...... Louis Trichardt Wh ..... Windhoek Lii . Liideritzbucht Wi ...... Willowmore WI ...... Williston Ma ...... Maseru Wo .. Worcester Mi ...... Middelburg Contributorsl

C. Bruckmann PD Dr. H. E. K. Hartmann Hamburger Berg 25 Institut fUr Allgemeine Botanik 20359 Hamburg OhnhorststralSe 18 Germany 22609 Hamburg Germany P. Chesselet Compton Herbarium Prof. Dr. H.-D. Ihlenfeldt Kirstenbosch Research Centre Institut fUr Allgemeine Botanik Private Bag X7 OhnhorststralSe 18 Claremont 7735 22609 Hamburg South Africa (RSA) Germany Prof. D. T. & N. A. Cole C. Klak P.o. Box 447 Bolus Herbarium Lonehill 2062 University of Cape Town South Africa (RSA) Rondebosch 7700 South Africa (RSA) Dr. M. Gerbaulet Phyllobolic Institute I. Niesler Friedrich-Soltau-StralSe 13 Institut fUr Allgemeine Botanik 29456 Hitzacker OhnhorststralSe 18 Germany 22609 Hamburg Germany L. B. Groen Paleis Het des Dr. S. M. Pierce National Museum Bolus Herbarium Koninklijk Park 1 University of Cape Town 7315 JA Apeldoorn Rondebosch 7700 The Netherlands South Africa (RSA) S.A. Hammer Dr. M. Struck-Gerbaulet 845 Mason Road KochstralSe 4 Vista California 92084 29451 Dannenberg U.S.A. Germany B. Hansen Dr. N. Zimmermann Lokstedter Steindamm 74 Friedrich-Engeis-StralSe 45c 22529 Hamburg 07749 Jena Germany Germany

Contributors of Photographs and Artist

M. B. Bayer, Paarl, South Africa; V. Bittrich, Campinas, South Africa; R. Kabutz, Pretoria, South Africa; C. Klak, Brazil; C. Bruckmann, Hamburg, Germany; R. Chin• Cape Town, South Africa; S. Liede, Bayreuth, Germany; nock, Adelaide, Australia; D. Cole, Lonehill, South Af• U. Meve, Bayreuth, Germany; 1. Niesler, Hamburg, Ger• rica; M. Dillon, Chicago, USA; M. Gerbaulet, Hitz• many; S. Pierce, Cape Town, South Africa; U. Schmie• acker, Germany; B. Groen, Wageningen, Netherlands; del, Hamburg, Germany; M. Struck, Dannenberg, Ger• H. E. K. Hartmann, Hamburg, Germany; H.-D. Ihlen• many; H. Toelken, Adelaide, Australia; C. Taylor, Mis• feldt, Hamburg, Germany; E. van Jaarsverld, Cape Town, souri, USA; J. Traeger, San Francisco, USA; R. Uijs, Pre• toria, South Africa; N. Zimmermann, lena, Germany. B. Braeutigam, Hamburg, Germany, has prepared 1 ••• of both volumes (A-E and F-Z). the ink versions of all drawings.