Aizoaceae (Illustrated Handbook of Succulent Plants) [2nd ed. 2017] 366249258X, 9783662492581

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Illustrated Handbook of Succulent Plants

Heidrun E.K. Hartmann Editor

Aizoaceae Second Edition

Illustrated Handbook of Succulent Plants Series Editors Urs Eggli Heidrun E. K. Hartmann

During evolution, many species have adapted to dry and arid environments by developing the ability to store water in their stems, leaves, or underground organs. These succulents can be found in a wide range of plant families. The Illustrated Handbook of Succulent Plants, consisting of five volumes, covers over 9000 taxa of succulents (excluding cacti). In addition to the volumes on Monocotyledons and Dicotyledons, separate volumes are devoted to those families with predominantly succulent members, which show an especially great diversity, namely Aizoaceae, Asclepiadaceae and Crassulaceae. Following an alphabetical listing of families, genera and species, detailed descriptions are given, including the taxonomy with synonyms, data on the distribution and ecology, references, and keys to genera, species or subspecies. Over 2000 superb colour photographs complete this inventory of succulent plants. More information about this series at https://link.springer.com/bookseries/4452

Heidrun E. K. Hartmann Editor

Aizoaceae Second Edition

With 694 Figures

Editor Heidrun E. K. Hartmann FB Biologie, Biozentrum Klein Flottbeck und Botanischer Garten Universität Hamburg Hamburg, Germany

ISBN 978-3-662-49258-1 ISBN 978-3-662-49260-4 (eBook) ISBN 978-3-662-49259-8 (print and electronic bundle) DOI 10.1007/978-3-662-49260-4 Library of Congress Control Number: 2017940217 # Springer-Verlag GmbH Germany 2002, 2017 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. Cover illustrations: Phyllobolus humilis (copyright M. Struck), Drosanthemum flammeum (copyright M.B. Bayer), Trianthema argentina (copyright R. Kiesling) Printed on acid-free paper This Springer imprint is published by Springer Nature The registered company is Springer-Verlag GmbH Germany The registered company address is: Heidelberger Platz 3, 14197 Berlin, Germany

Preface

About 15 years after the first edition of the handbook of Aizoaceae, the iceplant family, a wealth of changes can again be reported for the family, and this second edition brings a completely new survey over all taxonomic units from subfamilies down to species and subspecies; each critically examined. For all groups of users, e.g., farmers valuing the plants for their browsing stock, hobbyists loving to grow and flower mainly the “stone plants” of the family, and tourists enjoying the mass flowering of millions of annuals after good rains in the south-western part of South Africa, this book will offer again some improvements. But it will be of special interest to taxonomists, who still show some reservation against members of the Aizoaceae because of the size of the family and the confused and entangled systematics of many groups. To produce this handbook, the worldwide web knitted among hundreds of interested people for the first edition has been closer meshed and reinforced. 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 fieldwork – the last one by the editor in 2014, 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 having been made. As a result of all united efforts, an up-to-date survey of the state of the art can be presented, forming the base 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 Lexicon of Succulent Plants, published between 1970 and 1978 in different editions in German and English, in which Jacobsen treated 122 genera of Mesembryanthema under the heading “Mesembryanthemaceae.” In the two volumes of the first edition of this handbook dedicated to the family Aizoaceae, the Mesembryanthema were included. A total of 234 genera had been named in the family Aizoaceae, all of which were described, but only 136 could be accepted as valid. Thirty new combinations and two new names and two new species were published in the first edition; 189 taxa were lectotypified, 26 were neotypified. This second edition presents 146 genera in which the 1594 accepted of the presently known 2956 species are described. Seven genera have been newly v

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described since 2001, namely Brianhuntleya, Hammeria, Knersia, Malotigena, Phiambolia, Ruschiella, and Volkeranthus, and only two have been placed newly into synonymy (Caulipsilon and Muiria). All synonymous genus names are treated with the valid genus names. Results from preliminary molecular studies, using different groups of markers at different levels like species, genera, up to the subdivision of the family, which have influenced the taxonomy, are incorporated. The basic division into four subfamilies is supported by more recent studies, yet with some genera being shifted. Hundreds of new photos, most of them taken in habitat, illustrate the present situation. Furthermore, the geographical distribution of each genus is shown on a separate map – each revised according to new data – either worldwide or restricted to Southern Africa. For these latter, the main rainfall seasons are delimited by blue lines in order to offer also parameters for cultivation. The manuscript was revised and updated in the year 2014, the closing date for inclusion of known publications being 31 December 2014. I am most grateful to all authors and colleagues who have helped me to keep up with the numerous new descriptions, including contributions of taxa in the process of description and unpublished information. I thank all collaborators and contributors of text or artwork. My special thanks go to Dr. Lindenborn and to Susanne Friedrichsen, Springer Verlag, who made the publication possible at last. Hamburg, April 2016

H. E. K. Hartmann

While waiting for the proofs, my wife died on 11 July 2016, 5 days before the first chapters reached us. I am not a botanist, but having assisted my wife with her work in technical and administrative matters and knowing most of the authors, I considered it my responsibility to bring her work to a successful conclusion. With the help of the authors and Ms Susanne Friedrichsen, Senior Editor for Major Reference Works, I dared to face the challenge. While being capable of dealing with all printing and formatting issues, any decisions with regard to scientific issues were beyond my expertise, but for many problems I could find a solution in the files and papers of my late wife. In addition, I am particularly grateful to Dr. Maike Gerbaulet, who kindly replied to my request for scientific help and gave the maximum possible assistance under these sad circumstances. While we tried our best to eradicate all mistakes, errors, and inconsistencies, it is likely that some remain, which my wife would have spotted, had she had the opportunity. Sadly, she did not. Hamburg, October 2016

Wilfried Hartmann

Acknowledgements

Matt Buys thanks Cornelia Klak and Muthama Muasya. Steven Hammer and Andrew J. Young thank Chris Rodgerson, Terry Smale, Anthony Mitchell, Adam Harrower, and Nick Helme for their helpful and informative discussions on Conophytum and for their company during explorations in the veld, as well as Catherine Arthur, and Irma Burger. Their thanks go also to Danielle Rudeen of the Huntington Botanical Garden for endless xeroxing. They give particular thanks to the late Heidrun Hartmann: “amicitia vera illuminat.” H.E.K. Hartmann is most grateful to more than 700 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 43 years of studies in Aizoaceae including fieldwork in Argentina, Botswana, Ethiopia, Ghana, India, Kenya, Lesotho, Mexico, Namibia, South Africa, Sudan, Swaziland, Tanzania, UAR, the 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, botanical gardens, or military grounds • Getting permits from nature conservation bodies for collecting protected plants in all countries visited • Organizing sometimes extensive field trips • Allowing to stay overnight and feeding her well • Searching for plants in sometimes rather inaccessible places and under difficult conditions • Collecting data on distribution • Taking her to places, keeping her company and helping during fieldwork, 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 drawing leaves, fruits, and flowers • Discussing problems in finding roads or drawing the correct taxonomic conclusions, and, above all, being willing to share their knowledge with her

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The directors of the herbaria B, BM, BOL, BOLO, HEL, HBG, K, NBG, PRE, and 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 especially acknowledged for very extensive loans over a much extended period. Financial support for the work resulting in the first edition from the German Science Foundation (DFG) since 1969, from the Council of Scientific and Industrial Research in South Africa, the award of the Kirstenbosch Jubilee Fund 1977, from the National Botanic Gardens via the Karoo Botanic Garden, the Universities of Cape Town and Pretoria, the Hansische Universitätsstiftung of the University of Hamburg, the University of Hamburg, and the Cactus and Succulent Society of America is still most gratefully acknowledged. I.M. Niesler thanks Katrin Baumgärtner for her intensive care for the collection.

Acknowledgements

Contents

Volume 1 Part I

The Family Aizoaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1

Aizoaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. E. K. Hartmann and M. Gerbaulet

3

Part II

Genera of the Family Aizoaceae

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23

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25

Acrosanthes AIZOOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. E. K. Hartmann

31

Aethephyllum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. E. K. Hartmann

37

Aizoanthemum AIZOOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. E. K. Hartmann

41

Aizoon AIZOOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. E. K. Hartmann

45

Aloinopsis RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. E. K. Hartmann

53

Amphibolia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. E. K. Hartmann

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69

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73

Acrodon RUSCHIOIDEAE H. E. K. Hartmann

Antegibbaeum RUSCHIOIDEAE H. E. K. Hartmann Antimima RUSCHIOIDEAE H. E. K. Hartmann Apatesia RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113

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Aptenia MESEMBRYANTHEMOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 117 M. Gerbaulet Arenifera RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123 H. E. K. Hartmann Argyroderma RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127 H. E. K. Hartmann Aridaria MESEMBRYANTHEMOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 137 M. Gerbaulet Aspazoma MESEMBRYANTHEMOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . 149 M. Gerbaulet Astridia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153 H. E. K. Hartmann Bergeranthus RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161

Bijlia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169 H. E. K. Hartmann Braunsia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173 H. E. K. Hartmann Brianhuntleya RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179 H. E. K. Hartmann Brownanthus MESEMBRYANTHEMOIDEAE . . . . . . . . . . . . . . . . . . . . . . . 185 M. Gerbaulet Calamophyllum RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197

Callistigma MESEMBRYANTHEMOIDEAE M. Gerbaulet

. . . . . . . . . . . . . . . . . . . . . . . . 199

Carpanthea RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203 H. E. K. Hartmann Carpobrotus RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207 H. E. K. Hartmann Carruanthus RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215 H. E. K. Hartmann Caryotophora RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219 H. E. K. Hartmann Cephalophyllum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221 H. E. K. Hartmann Cerochlamys RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241 H. E. K. Hartmann

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Chasmatophyllum RUSCHIOIDEAE H. E. K. Hartmann

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Cheiridopsis RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251 H. E. K. Hartmann Circandra RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 273 H. E. K. Hartmann Cleretum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275 H. E. K. Hartmann Conicosia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281 H. E. K. Hartmann Conophytum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287 S. A. Hammer and A. J. Young Corpuscularia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 361 H. E. K. Hartmann Cryophytum MESEMBRYANTHEMOIDEAE . . . . . . . . . . . . . . . . . . . . . . . 369 M. Gerbaulet Cylindrophyllum RUSCHIOIDEAE H. E. K. Hartmann

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Cypselea SESUVIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383 H. E. K. Hartmann Dactylopsis MESEMBRYANTHEMOIDEAE M. Gerbaulet

. . . . . . . . . . . . . . . . . . . . . . . . 387

Deilanthe RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 391 H. E. K. Hartmann Delosperma RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 395 H. E. K. Hartmann Dicrocaulon RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 445 H.-D. Ihlenfeldt Didymaotus RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 453 H. E. K. Hartmann Dinteranthus RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 457 H. E. K. Hartmann Diplosoma RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 463 H.-D. Ihlenfeldt Disphyma RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 469 H. E. K. Hartmann Dorotheanthus RUSCHIOIDEAE H. E. K. Hartmann

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Dracophilus RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 483 H. E. K. Hartmann Drosanthemum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 487 H. E. K. Hartmann Eberlanzia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 541 H. E. K. Hartmann Ebracteola RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 549 H. E. K. Hartmann Ectotropis RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 555 H. E. K. Hartmann Enarganthe RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 559 H. E. K. Hartmann Erepsia RUSCHIOIDEAE H. E. K. Hartmann

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Esterhuysenia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 579 E. van Jaarsveld Eurystigma MESEMBRYANTHEMOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . 587 M. Gerbaulet Faucaria RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 591 H. E. K. Hartmann Fenestraria RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 599 H. E. K. Hartmann Frithia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 603 H. E. K. Hartmann Galenia AIZOOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 607 H. E. K. Hartmann Gibbaeum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 619 J. Thiede Glottiphyllum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 641 H. E. K. Hartmann Gunniopsis AIZOOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 653 H. E. K. Hartmann

Volume 2 Hallianthus RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 659 H. E. K. Hartmann Hammeria RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 663 H. E. K. Hartmann

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Hartmanthus RUSCHIOIDEAE H. E. K. Hartmann

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Hereroa RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 671 H. E. K. Hartmann Hymenogyne RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 681 H. E. K. Hartmann Ihlenfeldtia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 685 H. E. K. Hartmann Jacobsenia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 689 H.-D. Ihlenfeldt Jensenobotrya RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 695 H. E. K. Hartmann Jordaaniella RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 697

Juttadinteria RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 703 H. E. K. Hartmann Khadia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 709 H. E. K. Hartmann Knersia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 715 H. E. K. Hartmann Lampranthus RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 719 H. E. K. Hartmann Lapidaria RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 763

Leipoldtia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 767 H. E. K. Hartmann Lithops RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 777 D. T. Cole and N. A. Cole Machairophyllum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 811 H. E. K. Hartmann Malephora RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 815

Malotigena RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 825 H. E. K. Hartmann Marlothistella RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 829 H. E. K. Hartmann Mesembryanthemum MESEMBRYANTHEMOIDEAE . . . . . . . . . . . . . . . . 833 M. Gerbaulet and H. E. K. Hartmann

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Mestoklema RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 861 H. E. K. Hartmann Meyerophytum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 865 H.-D. Ihlenfeldt Mitrophyllum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 869 H.-H. Poppendieck Monilaria RUSCHIOIDEAE H.-D. Ihlenfeldt

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 879

Mossia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 885 H. E. K. Hartmann Namaquanthus RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 889 H. E. K. Hartmann Namibia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 891 H. E. K. Hartmann Nananthus RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 895 H. E. K. Hartmann Nelia RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 903

Neohenricia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 907 H. E. K. Hartmann Octopoma RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 911 H. E. K. Hartmann Odontophorus RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 917 H. E. K. Hartmann Oophytum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 923 H. D. Ihlenfeldt Opophytum MESEMBRYANTHEMACEAE M. Gerbaulet Orthopterum RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . 927

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 933

Oscularia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 937 M. H. Buys Ottosonderia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 945 H. E. K. Hartmann Peersia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 949 H. E. K. Hartmann Pherolobus RUSCHIOIDEAE H. E. K. Hartmann

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Phiambolia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 957 H. E. K. Hartmann Phyllobolus MESEMBRYANTHEMOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . 963 M. Gerbaulet Pleiospilos RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 991 H. E. K. Hartmann Plinthus AIZOOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 999 H. E. K. Hartmann Polymita RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1003 H. E. K. Hartmann Prenia MESEMBRYANTHEMOIDEAE M. Gerbaulet Prepodesma RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . 1007

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1017

Psammophora RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1021 H. E. K. Hartmann Psilocaulon MESEMBRYANTHEMOIDEAE M. Gerbaulet

. . . . . . . . . . . . . . . . . . . . . . . . 1027

Rabiea RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1041 H. E. K. Hartmann Rhinephyllum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1045 H. E. K. Hartmann Rhombophyllum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1051 H. E. K. Hartmann Ruschia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1057 H. E. K. Hartmann Ruschianthemum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1115 H. E. K. Hartmann Ruschianthus RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1119

Ruschiella RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1121 H. E. K. Hartmann Saphesia RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1125

Sarcozona RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1127 H. E. K. Hartmann Sceletium MESEMBRYANTHEMOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 1131 M. Gerbaulet

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Schlechteranthus RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . 1141

Schwantesia RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1145

Scopelogena RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1153

Sesuvium SESUVIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1157 H. E. K. Hartmann Skiatophytum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1169 H. E. K. Hartmann Smicrostigma RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1173

Stayneria RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1177 H. E. K. Hartmann Stoeberia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1179 H. E. K. Hartmann Stomatium RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1185 H. E. K. Hartmann Synaptophyllum MESEMBRYANTHEMOIDEAE . . . . . . . . . . . . . . . . . . . . 1197 M. Gerbaulet Tanquana RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1201 H. E. K. Hartmann Tetragonia AIZOOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1205

Titanopsis RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1225 H. E. K. Hartmann Trianthema SESUVIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1231

Tribulocarpus SESUVIOIDEAE J. Thiede

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1253

Trichodiadema RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1257 I.M. Niesler Vanheerdea RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1277 H. E. K. Hartmann Vanzijlia RUSCHIOIDEAE H. E. K. Hartmann

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1281

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Vlokia RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1285 H. E. K. Hartmann Volkeranthus MESEMBRYANTHEMOIDEAE . . . . . . . . . . . . . . . . . . . . . . . 1289 M. Gerbaulet Wooleya RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1295 H. E. K. Hartmann Zaleya SESUVIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1299 H. E. K. Hartmann Zeuktophyllum RUSCHIOIDEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1305 H. E. K. Hartmann List of All Generic Names in Alphabetical Order . . . . . . . . . . . . . . 1309

List of Abbreviations and Symbols

Ø  >  <   ! =

 § App. Art. b c. Chr comb. nov. Distr Ecol Etym f Fl Fr Gk h hort. I ICBN ICN

Diameter More or less Bigger than Bigger than or equal to Smaller than Smaller than or equal to Times; hybrid sign in a name The cited herbarium sheet has been seen Equals; between names: heterotypic synonym, based on a different type than the accepted name of the taxon Homotypic synonym, based on the same type as the accepted and valid name Section Appendix Article Broad Circa, about Chromosomes Combinatio nova, new combination Distribution Ecology Etymology Figure Flower(s) Fruit(s) Greek High Hortus, garden Internode(s) International Code of Botanical Nomenclature International Code of Nomenclature

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List of Abbreviations and Symbols

in litt. Inf ISO K l L La LT nomen illeg.

In litteris, in a letter Inflorescence(s) Isotype(s) Calyx Long Leaf, leaves Latin Lectotype Nomen illegitimum, illegitimate name, not permitted under the ICBN A or ICN@ nomen invalid. Nomen invalidum, invalid name, not fulfilling the requirements for valid publication as given by ICBN A or ICN@ nov. Nova, novum, new NT Neotype p.a. Per annum, in a year pers. comm. Personal communication p.p. Pro parte, in part S Seed(s) s.lat. Sensu latiore, in the wider sense s.n. Sine numero, without a number ssp. Subspecies s.str. Sensu stricto, in the narrow sense subg. Subgenus t. Tabula, plate t Thick T Holotype var. Varietas, variety 2n Double chromosome set

Abbreviations of names of authors of plant names follows Brummitt R.K. & Powell C.E. (1992) Authors of Plant Names. Royal Botanic Gardens. Kew in many cases, e.g. Haw. for Haworth, L. for Linnaeus, or N.E.Br. for N.E. Brown. Deviations have been found useful in cases in which the authors are rare within the family. Abbreviations of names of herbaria follows Holmgren et al. (1990) Index Herbariorum, Ed. 8 Names of genera are abbreviated by their first, capital, letter within the text covering that genus. An exception has been made for M, in which case M. always stand for the genus Mesembryanthemum; all other genera starting with an M are abbreviated by their first two letters, e.g. Ma. for Malephora or Machairophyllum, respectively.

Abbreviations of Geographical Terms Ab A.B. Ag Au B.E. Bi Bl Br Bu B.W. Ca Cl Co Cr C.T. E EC E.L. Fr FS Ga GAU Gb Ge Gh Gr G.R. Ha Ho Ja Jo Ke Ki K.K. KN Ku La L.B. Li Lo L.T. Lü Ma

Aberdeen Alexander Bay Aggenys Aus Barkly East Bitterfontein Bloemfontein Bredasdorp Burgersdorp Beaufort West Carnarvon Calvinia Carolina Cradock Cape Town East, eastern Eastern Cape Province East London Fraserburg Free State, Province Garies Gauteng, Province Gaborone George Grahamstown Griquatown Graaff Reinet Harrismith Hondeklipbaai Jagersfontein Johannesburg Kenhardt Kimberley Kap Kruis KwazuluNatal, Province Kuruman Laingsburg Lamberts Bay Limpopo Loeriesfontein Louis Trichardt Lüderitzbucht Maseru xxi

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Mi Mo MM MP Mu N NC No NP NW OR Ou P.A. Pc P.E. Pi Pm P.N. Po Pr Rc Ri Ru S S Africa Sp St Su Sw To Ui Um Un Up Va Vr VR W Wa WC Wh Wi Wl Wo

Abbreviations of Geographical Terms

Middelburg Montagu Mmabatho Mpumalanga, Province Murraysburg North, northern Northern Cape Province Noorduwer Northern Province North Western Province Orange River Oudtshoorn Prince Albert Potchefstroom Port Elizabeth Pietersburg Posmasburg Port Nolloth Pofadder Pretoria Richmond Riversdale Rustenburg South, southern Republic of South Africa Springbok Steytlerville Sutherland Swakopmund Touwsrivier Uitenhage Umtata Uniondale Upington Vanrhynsdorp Vryburg Vaal River West, western Warmbad Western Cape Province Windhoek Willowmore Williston Worcester

Abbreviations of Place Names in the Maps

Ab A.B.. Ag Au B.E. Bi Bl Br Bu B.W. Ca Cl Co Cr C.T. E.L. Fr Ga Gb Ge Gh Gr G.R. Ha Ho Ja Jo Ke Ki K.K. Ku K.K. La L.B. Li Lo

Aberdeen Alexander Bay Aggenys Aus Barkly East Bitterfontein Bloemfontein Bredasdorp Burgersdorp Beaufort West Carnarvon Calvinia Carolina Cradock Cape Town East London Fraserburg Garies Gaborone George Grahamstown Griquatown Graaff Reinet Harrismith Hondeklipbaai Jagersfontein Johannesburg Kenhardt Kimberley Kap Kruis Kuruman Kap Kruis Laingsburg Lamberts Bay Limpopo Loeriesfontein

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L.T. Lü Ma Mi Mo MM Mu No OR Ou P.A. P.E. Pc Pi Pm P.N. Po Pr Rc Ri Ru Sp St Su Sw To Ui Um Un Up Va Vr VR Wa Wh Wi Wl Wo

Abbreviations of Place Names in the Maps

Louis Trichardt Lüderitzbucht Maseru Middelburg Montagu Mmabatho Murraysburg Noorduwer Orange River Oudtshoorn Prince Albert Port Elizabeth Potchefstroom Pietersburg Posmasburg Port Nolloth Pofadder Pretoria Richmond Riversdale Rustenburg Springbok Steytlerville Sutherland Swakopmund Touwsrivier Uitenhage Umtata Uniondale Upington Vanrhynsdorp Vryburg Vaal River Warmbad Windhoek Willowmore Williston Worcester

Frequently Used References

ABH = Ann. Bolus Herb. = The Annals of the Bolus Herbarium 1–4. Cambridge University Press. London. 1914–1928. ASAM = Ann. S. African Mus. = Annals of the South African Museum BB = Bibliothec. Bot. = Bibliotheca Botanica. E. Schweizerbart’sche Verlagsbuchhandlung. Stuttgart. BBP = Beitr. Biol. Pflanzen = Beiträge zur Biologie der Pflanzen BCSJ = Brit. Cact. Succ. J. =British Cactus and Succulent Journal BJS = Bot. Jahrb. Syst. = Botanische Jahrbücher für Systematik BSKP = Beitr. Sukkulentenk. Sukkulentenpflege = Beiträge zur Sukkulentenkunde und -pflege CBH = Contr. Bolus Herb. = Contributions from the Bolus Herbarium CBM = Curtis’s Bot. Mag. = Curtis’s Botanical Magazine CJ = Cact. J. = Cactus Journal (Croydon) CSJ = Cact. Succ. J. (US) = Cactus and Succulent Journal (U.S.) CSJGB = Cact. Succ. J. Gr. Brit. = Cactus and Succulent Journal of Great Britain FPSA = Fl. Pl. South Africa = Flowering Plants of South Africa FR = Feddes Repert. = Repertorium specierum novarum regni vegetabilis GC = Gard. Chron. = Gardeners’ Chronicle ser.3 GF = Gartenflora = Gartenflora GW = Gartenwelt = Gartenwelt ICBN = International Code of Botanical Nomenclature (see Greuter et al. and McNeill et al.) ICN = International Code of Nomenclature JB = J. Bot. = Journal of Botany JSAB = J. S. African Bot. = Journal of South African Botany KB = Kew Bull. = Kew Bulletin of Miscellaneous Information KuaS = Kakt. And. Sukk. = Kakteen und andere Sukkulenten MBMUZ = Mitt. Bot. Mus. Univ. Zürich = Mitteilungen aus dem Botanischen Museum der Universität Zürich MBSM = Mitt. Bot. Staatssamml. München = Mitteilungen der Botanischen Staatssammlung München MDG = Möll. Deutsche Gärtnerz. = Möllers Deutsche Gärtnerzeitung

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Frequently Used References

MDK = Monatsschr. Deutsche Kakt.-Ges. = Monatsschrift der Deutschen Kakteengesellschaft MIBH = Mitt. Inst. Allg. Bot. Hamburg = Mitteilungen aus dem Institut für Allgemeine Botanik Hamburg MKK = Monatsschr. Kakteenkunde = Monatsschrift für Kakteenkunde NCSJ = Natl. Cact. Succ. J. = The National Cactus and Succulent Journal NM = Notes Mesembr. 1–2 = Notes on Mesembrianthemum and Allied Genera 2. 1928–1935. University of Cape Town, Cape Town. NM = Notes Mesembr. 3 = Notes on Mesembryanthemum and Allied Genera 3. 1936–1958. University of Cape Town, Cape Town. SAG = S. African Gard. = South African Gardening and Country Life SAJB = S. African J. Bot. = South African Journal of Botany SKK = Sukkulentenkunde = Sukkulentenkunde. Jahrbuch der Schweizer Kakteengesellschaft SUC = Succulenta (Netherlands) = Succulenta. Nederlands-Belgische Vereniging van Liefhebbers van Cactussen en ander Vetplanten. TRSSA = Trans. Roy. Soc. South Africa = Transactions of the Royal Society of South Africa VNVH = Verh. Naturwiss. Vereins Hamburg = Verhandlungen des naturwissenschaftlichen Vereins Hamburg (NF) ZSK = Z. Sukkulentenk. = Zeitschrift für Sukkulentenkunde

About the Editor

Heidrun (Heidi) E. K. Hartmann (1942–2016) Heidrun Hartmann wrote her M.Sc. thesis on the genus Harpagophytum in 1968 (published in 1970), which has thick tubers used also for medicine, but she considered her start into the world of succulence from 1969, when on July 19th she left for South Africa to begin her investigations of the genus Argyroderma in the Knersvlakte under the supervision of Prof. H.-D. Ihlenfeldt. She repeated her exercises to walk the area for another 4 months in 1971 and finished her Ph.D. in 1973: a revision of the genus reducing the 100 species to 10 (published in 1978). In 1974, she expanded her fieldwork to all genera with many locules in their capsules, with a focus on the subtribe Leipoldtiinae, on the taxonomy of which she wrote my habilitation thesis handed in 1982. In collaboration with several of her students at the University of Hamburg, she has published a number of revisions of most genera included in the subtribe until her death, plus some others like that of Glottiphyllum or Pleiospilos. Fieldwork has been the major source of her investigations, and her interest in other areas of Africa was raised early. Since 1969, she has worked in Botswana, Ethiopia, Ghana, Kenya, Lesotho, Malawi, Namibia, South Africa, Sudan, Swaziland, Tanzania, and Zimbabwe; for comparison of habitats and growth forms, she has visited Argentina, Mexico, and the southwestern states of USA; In 2007, she has undertaken investigations in the United Arab Emirates.

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Since 1970, she was employed as a lecturer at the Botanical Institute of the University of Hamburg and since 1982 in the function of a professor. Beside supervising numerous students in her field of systematic botany, this task implied, e.g., also more than two decades as head of the Ph.D. Commission of the Faculty guiding more than 80 students through their final phase of the Ph.D. process. Although she enjoyed teaching very much, especially lecturing on new subjects, as during her last years, she was happy being retired since October 1, 2007, after having turned 65 in August that year, and she remained a member of the university. Her scientific work at the University of Hamburg until her death was supported by cultivation of material in a ca. 600 m2 greenhouse of the Botanical Garden of the University of Hamburg, diligently supervised by Katrin Baumgärtner since many years. Since 1969, Heidi has taken black-and-white photographs of material she collected as well as slides, the collection amounting to about 45,000 items, including electronic images since 2005. Another important tool was also a data bank of almost 15,000 sets of data in ACCESS, into which she had entered all collections she had made, alone or with students or colleagues of hers. She had been able to buy a program “Taxo2map” to work from this databank to develop distribution maps that are precise to the second and can, at the same time, be altered if need be, e.g., in cases of name changes. This combined application allowed a very fast examination of additional data, e.g., from other collections and from type material of older species, to check exact areas and probabilities of correct identifications and to publish the resulting maps. Since her retirement, she was concentrating on four topics: 1. Molecular and classical morphological work on the weedy Aizoaceous genus Trianthema together with Sigrid Liede-Schumann, professor at Bayreuth and her first PhD student once. A first paper based on extensive morphological studies is available since June 2011 (Hartmann, Meve and Liede-Schumann in Plant Ecology and Evolution) 2. Studies on nectaries in Aizoaceae together with Dr. Ingeborg Niesler who has worked mainly on Trichodiadema and leaf surfaces, being an expert in SEM work. Nectaries and leaf surfaces are of uttermost importance for a better understanding of the evolution of the Mesembryanthema since character states of the “other Aizoaceae” reveal, e.g., in nectaries, the basal conditions from which features found higher up in the tree of evolution have derived (Hartmann and Niesler 2009). 3. Molecular and classical morphological work on the genus Drosanthemum in collaboration with Sigrid Liede and Ulrich Meve as well as Anso le Roux, an ecologist in South Africa, and with students in MSc courses at Bayreuth since 2008. 4. Edition of the second edition of this Handbook of Aizoceae, including the rewriting of many of the genera included in the family, bringing the information up to date.

About the Editor

About the Editor

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After her retirement, she undertook studies in the genus Delosperma in northeastern Africa and southwestern Arabia (Hartmann 2008) and, published in a separate paper, in Gauteng (Hartmann 2009). Thanks to the Internet, she was in contact with researchers of Aizoaceae, including also ecologists, from Australia over the Near East and Kenya to South Africa, South America, and the USA as well as with colleagues in Europe. Exchange of data and information was intensive and most efficient, and she was very glad that she could make use of all these facilities at liberty from her home desk, given the fact that due to her heavy arthritis she had to rely on a wheel chair during her last 5 years. She drafted this text in October 2011. Since then, she has published 12 new articles and finished the rewriting and editing of the 2nd edition of this handbook. She received the “Cactus d’Or” in 2012 in recognition of her decades of work on succulents, especially Aizoaceae, and undertook the trip to receive it in 2013 on occasion of the “Expo Monaco 2013.” In 2014, she traveled to South Africa again and did field work with the help of friends though chairbound. But much to her disappointment, she was too weak to be present in person when she was awarded Fellowship of the CSSA in 2015. She died on July 11, 2016.

Contributors

Matt H. Buys New Zealand Forest Research Institute (Scion), Rotorua, New Zealand Desmond T. Cole Lonehill, South Africa Naureen A. Cole Lonehill, South Africa Maike Gerbaulet Phyllobolic Institute, Hitzacker, Germany Steven A. Hammer Vista, CA, USA Heidrun E. K. Hartmann Hamburg, Germany Biozentrum Klein Flottbek, Hamburg, Germany Hans-Dieter Ihlenfeldt Waabs, Germany Biozentrum Klein Flottbek, Hamburg, Germany Ernst J. van Jaarsveld Babylonstoren Farm, Simondium, Western Cape, South Africa Department of Biodiversity and Conservation, University of the W. Cape, Bellville, South Africa Ingeborg M. Niesler Biozentrum Klein Flottbek, Hamburg, Germany Hans-Helmut Germany

Poppendieck Biozentrum

Klein

Flottbek,

Hamburg,

Joachim Thiede Hamburg, Germany Andrew J. Young, Neston, Cheshire, UK Liverpool John Moores University, Liverpool, UK

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Contributors of Photographs and Artists

M.B. Bayer, Paarl, South Africa; C. Barnick, Hamburg, Germany; V. Bittrich, Campinas, Brazil; P.V. Bruyns, Rondebosch, South Africa; M.H. Buys, Rotorua, New Zealand; R. Chinnock, Adelaide, Australia; J. Chwekun, La Mesa, U.S.A.; D. T. Cole, Lonehill, South Africa; M. Gerbaulet, Hitzacker, Germany; H. Gölling, Harsum, Germany; S. Hammer, Vista, U.S.A; H.E.K. Hartmann, Hamburg, Germany; N.V. Helme, Scarborough, South Africa; H.-D. Ihlenfeldt, Waabs, Germany; E. van Jaarsveld, Kirstenbosch, South Africa; N. Jürgens, Hamburg, Germany; R. Koopman, Cape Town, South Africa; A. le Roux, Pretoria, South Africa; S. LiedeSchumann, Bayreuth, Germany; U. Meve, Bayreuth, Germany; U. Mierwald, Kiel, Germany; M. Mougin, Rieux-Minervois, France; I.M. Niesler, Hamburg, Germany; H.-H. Poppendieck, Hamburg, Germany; R. Riddles, Worcester, South Africa; D. Roth, Hamburg, Germany; U. Schmiedel, Hamburg, Germany; C. Schröder, Bayreuth; A.L. SchuetteVlok, Johannesburg, South Africa; D. Smith, Colchester, U.K.; M. Struck, Hitzacker, Germany; M. Thulin, Uppsala, Sweden; C. Walker, Milton Keynes, U.K.; A. J. Young, Neston, U.K.

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Introduction

The Family Aizoaceae Circumscription and Delimitation of the Family Aizoaceae As in the first edition, the plant family Aizoaceae Martynov nomen conservandum is understood here in the sense first presented by Bittrich and Hartmann (1988: 239–254): Aizoaceae s.str. Since the beginning of the millennium, however, molecular investigations have initiated a major change in perception, a revolution, that can be seen from two completely different viewpoints: Several molecular studies have confirmed the circumscription of the family (Klak et al. 2003 using chloroplast DNA, Thiede 2004 analyzing nuclear mitochondrial DNA), both resulting in permitting a division into four different subdivisions now, ranked as four subfamilies (overviews, e.g., in Klak 2010; Klak et al. 2013; Thiede 2014). Even further, in recent approaches, the groupings based on molecular studies are mostly interpreted as unquestionable conclusions regarding the taxonomic consequences. In fact, however, the investigated gene sections as such form only technically determined portions of gene sequences, and further characters and their expressions have to be studied in addition. In general, it has to be kept in mind that normal morphological characters reflect also the basal gene set with respect to such features, and it is always important to investigate a clearly delimited range of the taxonomic unit, i.e., here: the entire family Aizoaceae. This approach is exemplified by an extensive and thorough study of nectaries together with features of the flower construction (Hartmann & Niesler 2009), permitting a new interpretation of the nectary over the entire family in presenting different evolutionary lines, identifying the basal flower type with a sterile tepal-stamen tube rising above the plain basal nectary ring in Sesuvioideae, correlated in that subfamily with the possession of a pyxidium as the fruit in most of them, those of Tribulocarpus not having been studied in detail until now. In the remaining three subfamilies, nectaries appear at the upper end of the flower tube, in a flat ring (Aizooideae incl. Tetragonia) or as solitary sections, as in Mesembryanthemoideae. In some genera of the Ruschioideae, tubes of different origins are formed: a hypanthium or

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flower cup is developed in Glottiphyllum and an androecial tube in Conophytum, both raising the androecial elements, while the calyx elements sit basally and remain free. In both these cases, the flowers are tightly enclosed by subtending leaves, hindering them to open fully. In consequence, data, e.g., from members of Mesembryanthema cannot be transferred to the family Aizoaceae as a whole, because Mesembryanthema comprise two well-separated subfamilies, Mesembryanthemoideae and Ruschioideae, representing ends of evolutionary lines. Within the order Centrospermae, the Aizoaceae can morphologically best be recognized by a number of probable synapomorphies, i.e., features more or less exclusively present in the primitive members of the family and absent in other families of the order (Bittrich & Hartmann 1988; Bittrich 1990: 493–494). These are – – – – –

Possession of epidermal bladder cells Perigynous flowers Very long funicles Absence of wood rays in the normal and abnormal secondary xylem Possession of expanding tissue acting on a hygrochastic base

As discussed by Bittrich and Hartmann (1988) and Bittrich (1990), only the possession of epidermal bladder cells is completely unique for members of Aizoaceae within the Centrospermae (= Caryophyllales), and consequently, only this one feature is a true synapomorphy. For all other probable synapomorphies listed above, an occurrence in other families can also be shown. This does not render the list above invalid, but more investigations in other families are needed in order to determine the evolutionary direction of the character states.

Subdivision of the Aizoaceae The introduction of molecular studies in Aizoaceae led to the distinction of four subfamilies: Klak et al. (2003) used four different chloroplast markers, namely atpB-rbcl, psbA-trnH, rp16, and trnL-F, resulting in a shift of Tribulocarpus into the Sesuvioideae, and of Tetragonia appearing together with samples from Gunniopsis next to Aizooideae. In this way, the subfamily Tetragonoideae disappeared and four subfamilies remain at present in the Aizoaceae: Aizooideae, Mesembryanthemoideae, Ruschioideae, Sesuvioideae. This same arrangement was confirmed by Thiede (2004), who analyzed ITS, i.e., nuclear sequences of the ribosomal DNA. Several of these subfamilies have been considered to represent families of their own: Aizoaceae in a stricter sense than used here, Mesembryanthemaceae for the two subfamilies Mesembryanthemoideae and Ruschioideae together, Sesuviaceae, and Tetragoniaceae (tables in Bittrich & Hartmann 1988: 246, 247). The contents of the families varied greatly, hence older placements of genera must be checked and adapted according to recent results and treatments.

Introduction

Introduction

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The four subfamilies of the Aizoaceaecan be arranged in two groups. One group, with petaloid staminodes or petals, comprises the subfamilies Mesembryanthemoideae and Ruschioideae and is, as mentioned above, sometimes ranked as a family by the conserved name Mesembryanthemaceae Fenzl (Bittrich & Hartmann 1988; Bittrich 1990). In any case, these two subfamilies represent independent distal branches united by the possession of petals, n = 9, and valve wings. The remaining two subfamilies may be grouped together, too, if the basic chromosome number of n = 8 (or 7), the petaloid coloring of the inner side of the calyx lobes, or the possession of calcium oxalate druses in the cell lumina (in contrast to raphide bundles) can be shown to be synapomorphies for this group (for discussion, see Bittrich 1990). The Mesembryanthemoideae and Ruschioideae share the possession of hygrochastic expanding tissue on the valves with the former Aizooideae (with five locules), and with Gunniopsis, but Tetragonia lacks any such mechanical opening mechanism. Tetragonia, however, shares with Gunniopsis the possession of four locules, forming a group of its own below the rank of subfamily, and both genera are focussed on Australia. Bittrich (1990) had already suggested that the Sesuvioideae could be a sister-group to the remaining subfamilies, a position confirmed by Klak et al. (2003) and Thiede (2004). The subfamilies Mesembryanthemoideae and Ruschioideae together are named Mesembryanthema (Hartmann 1991), a group without a taxonomic rank. The subfamilies, as represented here, do certainly not meet equal standards as regards the knowledge of their character states, nor do they present groups of equal rank in a cladistic sense. They are, however, monophyletic in all molecular studies. Further subdivisions into tribes and subtribes, begun already by Schwantes (1947), were established further for “Mesembranthemaceae” in more extension by Chesselet, Smith, and van Wyk (2001, 2002, 2005), based on counts of nectaries in flowers of various members of Mesembryanthema. The Mesembryanthemoideae possess plain coilomorphic nectaries in central placentation, representing a basal condition in the family (Hartmann & Niesler 2009); in contrast, in the subfamily Ruschioideae, the formation of parietal placentation has changed the flower construction fundamentally: The former tube is prominently reduced to a fold on top of the inferior ovary, developing the nectary or nectaries at the upper edge, leaving no space for a further tube at all (Hartmann & Niesler 2009). Hence, at this stage in the evolution, the positions of the nectaries interact in number and shape with the expanding tissues at the base of the valves of the capsules: In areas with nectariferous tissue, no material is available to form firm abutments of the expanding tissue. In consequence, counts of nectary numbers on open flowers of Ruschioideae can only reflect the space available after the function of the opening valves has been organized in ontogeny (Hartmann & Niesler 2009). Further, loss of the rpoC1 intron in parts of the Ruschioideae separates Apatesieae and Dorotheantheae from the majority of Ruschieae including the genus Drosanthemum (Thiede, Schmidt & Rudolph 2007).

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Geographical Distribution of the Family Members of the Aizoaceae inhabit the tropics and subtropics worldwide between 40 north and south (Fig. 1). For each genus of worldwide occurrence, a map is provided in comparable format, on a purely geographical base. In the New World, the distribution area stretches from the southern states of USA over Central America and the Carribean Islands along the coasts of South America to Chile less than 40 S in the west and to Buenos Aires in the east. In Asia, the southern countries are inhabited by members of the Aizoaceae from Turkey to Malaysia and as far north as Afghanistan and China, albeit mainly as weeds. In Australia, New Zealand, and on the adjacent islands, members of the family are found in various habitats, and best known is probably the New Zealand spinach, Tetragonia tetragonoides, the only species of the family grown as a vegetable. In Africa, including the Cap Verdes, the Canary islands, and the Madeira islands, only the low-lying parts of the tropics around the Congo Basin are devoid of Aizoaceae. Only few species occur in Europe, e.g., in southern Spain and Italy, in perhaps natural stands. There is little doubt that various species and genera found in localities further north are secondary immigrants, e.g., in southern England and on the Canary Islands. Yet, it remains uncertain which taxa have immigrated when. Later immigration is also suspected for some of the species found in California and perhaps also in South America, exemplified by the genus Carpobrotus. It must be remembered, however, that several species of the genus Tetragonia occur naturally along the western coast, i.e., the Pacific 80°

60°

80° 60°

40°

40°

20°

20°

0°

0°

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20°

40°

40° 60°

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Fig. 1 Map of the world with the reported areas all presently known members of the family Aizoaceae

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K.K. Wh

Pi

February Jo Lü

Au

A.B.

March OR No

VR Ku

Up

Wa Ag

Po

Ke

Pm

VR

Gr

Ki

Summer

Sp

Ho

Ma

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Pc

Ja

Ga

Lo

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Ca

Rc

Cl

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Ab

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Mo Ri Br

Ou Un Ge

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Mi

Um

Cr

G.R.

St

Gh

E.L.

Ui P.E.

All seasons

Fig. 2 Map of Southern Africa with names of places following the most recently used names of districts

Ocean. In spite of the very large distribution area of the Aizoaceae, the centers of diversity lie close together in the south-western part of Africa. Latest counts on the basis of the genera accepted here show, e.g., for a square of 3030' (i.e. a quarter of a degree square), 36 different genera in the Gariep centre, i.e. in the lower part of the Orange or Gariep River. For Southern Africa, it is now even possible to show a map indicating the main rainfall seasons (Fig. 2) for each genus (except for Calamophyllum for which no material is available), in this way permitting to determine the watering preferences in cultivation for this genus.

Diversity and Ecology of the Family The extremes of the ecological niches inhabited by members of the family Aizoaceae are dry subtropical deserts, wet tropical coasts, and snow-covered subtropical mountains.

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The deserts receive less than 100 mm rainfall per year; some of them, however, are exposed to regular fog from the sea, increasing the overall available humidity. Rain falls either in winter, e.g., in the southern Namib and adjacent coastal areas, where, e.g., Fenestraria and Psammophora grow, or in summer, e.g., along the edges of the Sahara and into East Africa, where, e.g., species of Zaleya or Aizoon are found. Wet tropical coasts are inhabited by some members of the family all around the world on all continents. On rocks by the sea, exposed to salty spray, e.g., Delosperma napiforme and Sesuvium edmonstonei occur. In brackish marshy areas, often near mangrove vegetation, and exposed to tidal floods, Sesuvium portulacastrum can form huge stands. This species has been transferred from different tropical islands to, e.g., coasts of the United Arab Emirates on purpose, in order to support sand surfaces (Hartmann 2007a, b). Snow covers the high peaks of the Drakensberg Mountains in Lesotho for part of the year; some species of Delosperma grow under these conditions, some of them as endemics, e.g., Delosperma nubigenum, which can consequently also be grown outdoors in moderate climates if protected from too much humidity in winter. A new genus, Malotigena Niederle (2012a, b), has been described from these high peaks. The climate, in which the highest number of genera and species is naturally found, can be described as semi-arid with 100–400 mm precipitation per year, the rain falling mainly in winter. The annual amounts of rain can vary considerably, e.g., in one locality between 110 and 330 mm on a single farm near Fraserburg, and the intermediate drying of the soil represents an important factor in the life cycle. Under these conditions, members of the family form a major portion of the vegetation cover, in the number of species and in biomass. Aizoaceae are therefore important elements in farming management in winter rainfall areas and the adjacent regions in Southern Africa.

Importance of Characters for the Delimitation of Taxa Within the Aizoaceae Haworth (1795–1831) used mainly vegetative characters to delimitate species, following Linnaeus (1753, 1762), who referred to Dillenius’ drawings (1732) in many cases. Brown (1925: 212) was the first to recognize that similar or apparently identical growth forms and leaf shapes were correlated with different types of flowers and fruits. Since then, capsule morphology has played an increasingly important role in delimiting genera, based on a first general typification by Schwantes (1952). The number of genera with a uniform or rather uniform type of fruit has increased over the decades (e.g., Argyroderma, Hartmann 1978; Antimima, Dehn 1989), albeit also genera have been found recently, in which certain features of the capsule morphology differ from well-known fruit types, e.g., in Khadia (Chesselet & Hartmann 1995). The species of Khadia differ in the anatomy and morphology of the closing bodies, which, nevertheless, appear to result in similar function. Capsule morphology has, in spite of all insight, been studied only in a restricted number of cases in enough detail to permit conclusions, perhaps due to the complexity of the mechanism and morphology. Capsules play insofar an

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important role in the identification of taxa as they remain on the plant for several years in most cases and can, therefore, be easily found and used in most cases reliably over more than one season, in some cases like in Ruschia adorning the plants over a sequence of up to 4 years. Data on capsule morphology are therefore included in the descriptions wherever they have been available. Detailed information can be drawn from Schwantes (1952) and Hartmann (1988, 1991), both citing relevant literature. Drawings of several typical forms are presented as a survey, including names used in the text for descriptions in Fig. 3. As a new set of characters, nectaries have been studied in detail and in correlation with the shape of flowers and especially with the position they take in the different flower types (Hartmann & Niesler 2009). The basic flower type is perigynous with a long tube raised above the basal region in which nectary glands are positioned sunken into the plain epidermal tissue, found in the Sesuvioideae (Hartmann & Niesler 2009: Diagram 3). Such “plain nectaries” have been shifted up the tube in Aizooideae, i.e., Aizoon, Galenia, Gunniopsis, and in Tetragonia, the latter combined with a short indented tube and equipped with an inferior ovary; no single nectaries can be counted in such flowers, but the nectary glands sit in an irregular ring or band.

Technical Information on the Lexicon General Data An introduction to the family, keys to the subfamilies, and their descriptions are given first, followed by keys to the genera for each subfamily. The core part of the book consists of the alphabetical list of genera, each genus with its species listed alphabetically. Here, the current state of the taxonomic and systematic exploration of the family is presented in a consistent and informative manner. To this end, the contributors have worked over more than a decade, have traveled tens of thousands of miles to collect more data and take many photographs in the field, have seen and checked about 5000 references in literature and have grown thousands of plants under various conditions. Above all, the emphasis is on the correct taxonomic nomenclature and identification of the taxa, including all references to original publications.

Terminology for Descriptions Due to the fact that leaf succulence is a dominant feature in Aizoaceae, some special terms have been created to describe particular character states, e.g., of single leaves in those genera in which not more than one leaf pair is developed, exemplified best by the genus Lithops. The relevant terminology is given in the appropriate places; only some general terms are discussed here. “Spines” in Aizoaceae are always organs derived from stems, mostly from pedicels or peduncles (e.g., in Ruschia subg. Spinosae); “prickles” are remnants of organs of various origin and generally blunter than spines (e.g., in Drosanthemum); “spinules” are sharp, pointed, tiny organs, mostly at nodes

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Fig. 3 Fruits of Aizoaceae. (a–d, h–k): longitudinal sections, (e–g): views of open capsules from above. (a) Aizooideae. Hygrochastic capsule opening incompletely, dispersal by wash-out mechanism.- Aizoon canariense. (b–g) Ruschioideae. Hygrochastic capsules with expanding keels and sheets on the valves, increasing enclosure of seeds and correlated dispersal mechanisms (elaborated splash-cup types). (b, e) Parallel central expanding keels, covering membranes absent, the seeds are visible. Dispersal by wash-out or undirected simple splash mechanism, the broad rectangular wings form a wide cup when the valves are open so that the seeds are ejected together with water droplets by rain. Sometimes, seeds are retained in a basal pocket.- Stomatium spec. (c, f) Divergent expanding keels permit the formation of covering membranes from valvar roof endocarp plus some mesocarpal layers from underneath the broad expanding keels pulled out at the first opening of the capsule. Covering membranes thin and very flexible, dispersal by temporary deformation, not directed. Note the irregular shape of the valve wings tapering into awns distally from a broad base.Tanquana archeri. (d, g) Firm covering membranes of constant in shape irrespective of the contents of the locule, undulate in radial view with a ± pronounced recurving at the distal end touching the mostly large closing bodies and resulting in a complete blocking of the distal opening of the locule. Additional closing devices at the base of the recurving as rodlets, ledges, or bulges. Dorsal appendages on the exposed surface of the covering membranes stabilize the construction aiding directed dispersal. Dispersal by ejection of seeds through slits in ± proximal elevated parts of the covering membranes, optimal ballistic curves can be reached by the seeds. D: Antimima spec., acd = a closing rodlet here, (g): Odontophorus angustifolius, acd = a closing ledge here. (h) Mesembryanthemoideae. Hygrochastic capsule opening hygroscopically: the upper half of the septum expands when wet; false septa form broad valve wings; dispersal by rain as simple splash-cup or wash-out mechanisms.- Phyllobolus trichotomus. (i) Sesuvioideae. Capsule opening by removal of the top half of the capsule in dry conditions.- Sesuvium sesuvioides. (k) Tetragonioideae. Winged nut,

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and derived from the central bundle of a withered leaf or the triangle between leaf bases (e.g., in Drosanthemum). In some rare cases, leaves can turn into spinulate organs with age, too. The fate of the leaf can vary in Aizoaceae. The most widespread type of hard, xeromorphic leaves remains in shape during the dry season, shrinking to some degree, depending on drought conditions. On heterophyllous plants, one leaf pair remains regularly on the plant in a more or less green, assimilating condition, the other – lower and outer – pair forming a sheath enclosing the new growth of the following season (e.g., Mitrophyllum). This latter whole leaf pair or at least the free leaf tips are marcescent to various degrees (e.g., in Cheiridopsis, Antimima). In a number of homophyllous genera, the whole leaves can turn into a marcescent or hard, dry sheath, persisting during the dry season (e.g., Conophytum). Intermediate and even more variable conditions have been found in few other genera; in Dicrocaulon, Meyerophytum, Monilaria, Diplosoma, and Oophytum, the plants are sometimes termed “deciduous.” True deciduous conditions, i.e., the complete shedding of leaves at the end of the season, regularly by means of an abscission layer, is very rare (e.g., in Drosanthemum deciduus, or in Phyllobolus, both taxa named after this feature). Flowers in Aizoaceae possess an outer perianth, which is termed perigon in those cases, in which only stamens are developed in the subsequent inner rings; the elements of a perigon are named tepals. In most genera of Aizoaceae, however, the originally androecial primordia multiply, and the outer ones develop into petaloid organs. These showy elements are often called “petaloid staminodes,” because of their ontogeny, and named “petals” in this book, because they act ecologically as the attracting organs for the pollinators. Between stamens and petals, more or less filamentous elements can be found in many flowers. Irrespective of their width, these parts of the flower are termed “filamentous staminodes,” if they differ sufficiently from the petals. Readers will find that in a number of taxa a merging of petals into filamentous staminodes is described (e.g., in Mesembryanthemoideae), allowing a distinction of inner and outer elements but not a clear-cut borderline between the two categories. Stamens are distinguished from filamentous staminodes by the possession of anthers. In cases, in which filaments and filamentous staminodes resemble each other closely, it may be difficult to separate members of the two categories because anthers fall off in aging flowers. At the base of the stamens, nectaries of different shapes can be found as described above. The primitive forms consist of plain cells shaped like stomata above secretory tissue exuding the nectar, arranged in a ring in the

ä Fig. 3 (continued) dispersal anemochoric or perhaps hydrochoric.- Tetragonia decumbens. acd additional closing device at base of distal recurving of covering membrane, cb closing body, cm covering membrane, cmm margin of covering membrane of mesocarpal origin, ct corky, spongy tissue, da dorsal appendage at apex of perigone-lobe, dc subcentral appendage of covering membrane, dl dehiscence line, dr distal recurving of covering membrane, ek expanding keel, e expanding sheet, fs basal false septum, fw fruit wing, p persistent dry perigone-lobe, pl placenta, pm translucent margin of perigone-lobe, pp protrusion of perigone, s septum, sb spongy body of white, slightly cutinised tissue, separated distinctly from ct, sct sclerotic tissue of nut, sr septal rim, v valve, vr valve rim, vw valve wing

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tube of the perigon, found in Sesuvioideae and Aizooideae, as described above. Best known are, however, the two types prevailing in Mesembryanthema: coilomorphic nectaries are shell-shaped and characterize members of the Mesembryanthemoideae; lophomorphic nectaries are crestshaped, dominating in the Ruschioideae. Of the gynoecium, the stigmas are the parts most easily visible, representing the pollen-receiving free portions; styles, the connate lower portion of stigmas, occur very rarely in Aizoaceae (e.g., in Argyroderma). During anthesis, the flowers open and close by basal growth of all elements, resulting in an increase in diameter during the flowering period, a feature to be observed when measuring daily values. The flowers of Aizoaceae are protandrous, i.e., the male organs open first, hiding the gynoecium more or less completely. In most species of Delosperma in North-East Africa however, flowers are protogynous, i.e., the female phase in most flowers precedes the male (Hartmann, Liede 2013). In many flowers, the stigmas spread during the female phase, often elongating and spreading widely over the withered stamens. These developmental changes should be considered when floral characters are compared. Fruits of Aizoaceae are mostly hygrochastic capsules, very rarely berries (e.g., Carpobrotus), nuts (e.g., Tetragonia), or drupes (e.g., Gunniopsis). For the majority of the hygrochastic capsules of the family with many very special morphological components, an elaborate terminology has been developed. Terms used in this book are explained in Fig. 3 and, in addition, specified in cases of necessity in some genera (e.g., Caryotophora). Descriptions of fruit types follow those described by Hartmann (1988), descriptions are repeated here with those genera after which the type is named.

Taxonomic Principles Rank and Limits of Genera During the intensified investigations in the family in the past decades, the question of generic rank has been discussed several times (e.g., by Hartmann 1983: 46–48; Bittrich 1987:65–66; Hartmann BJS 1992: 43–46) suggesting criteria for the establishment of a taxon at genus rank. Many genera have been described in the early period of splitting up the old genus Mesembryanthemum based on one or few extraordinary character expressions: for example Fenestraria with club-shaped leaves with a terminal window, Amphibolia combining a closing body and broad valve wings in the fruit, or Esterhuysenia without valve wings and closing bodies in the fruit. In the absence of detailed investigations, several of these often monotypic genera have been reduced to synonymy of various big genera (e.g., Rowley 1978: 6–9, 62), enlarging these latter ones past recognition and definable limits. In the course of further studies, an overall genus concept will probably be developed, but until then a pragmatic view has to be taken. For the present treatment, the guideline has been to delimit genera as such when some to few clear distinctive character states can be found which, on one hand, permit a clear

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circumscription of the genus, and on the other hand allow a distinct separation from other genera. In most cases, this leads to traditional use, but in some cases, remarkable new arrangements had to be made. Thus, e.g., the genera Eberlanzia and Amphibolia can clearly be separated based on fruit and leaf character states, both genera can neither be united nor can they be placed in some other genus without unjustified distortions (Hartmann 1998a). On the same line of arguments, namely that taxa are recognized here as genera based on clear differences, the genera Peersia and Deilanthe have been re-established. Based on results of Dehn (1989: 189–216), the genus Antimima has been enlarged to about 100 species (Hartmann 1998b) that can probably be grouped into several subunits. For all these genera, a positive placement of the species included was possible, i.e., for all units below genus rank, the features found agree with the critical character expressions of the genus. In contrast to these re-established genera and their species, a number of genera remains containing several to many aberrant species. These species can, at this date, only be pointed out as being not in agreement with the established generic character states; at the same time, they cannot be placed elsewhere due to lack of data. The situation will be found particularly disturbing and dissatisfying in the large genera Delosperma, Drosanthemum, Lampranthus, and Ruschia, which need a revision urgently. In these, and in other cases where no recent results are available, a very conservative view has been taken keeping up most species in the genus into which they had originally been placed.

Rank and Limits of Species Of the numerous definitions of species, a simple one will serve the purpose of this handbook: species are biological units comprising all members of a group that can cross and produce viable offspring in habitat. As an approximation to this standard, which is not readily testable in perennial plants, a certain degree of similarity in all possible character expressions is considered sufficient to establish a species. Yet, even this level of treatment has seldom been reached. Instead, most species in Aizoaceae have been described based on a single specimen, available in the dry state as a herbarium specimen only, or as a growing plant. These latter have often been in cultivation for some period under conditions differing markedly from those of the natural habitat. In extreme cases, dry and cultivated plants have been described as different species. Early studies in habitat, including the variation in character states, have been extremely rare (e.g., Dinter 1923, 1928; Marloth 1910, 1914). This is particularly true for the period between 1920 and 1940, when most species were described by Bolus (Cape Town), Brown (London), and Schwantes (Kiel). Since for these descriptions no information regarding the variability was known, boundaries of species had necessarily to be uncertain, resulting in severe discrepancies about limits, affinities, and identities of taxa, and causing numerous discussions and hot debates among the main authors. The situation turned only when a revision of the genus Argyroderma was presented (Hartmann 1973, published 1978), which was based on extensive fieldwork including ecology and population biology, considering a wide array of characters, using statistical methods, and offering a species concept for succulent plants for the first time. It was not surprising that the number of species in

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Argyroderma was consequently reduced from 62 to 11, including one hybrid species. Later revisions of several bigger genera conducted in the same way, e.g., Cheiridopsis or Cephalophyllum, have reduced the number of species by about a third. Whether a similar effect will be found once the large genera, e.g., Delosperma, Lampranthus, and Ruschia have been studied, remains to be seen. The small genera with few species are mostly too well known to house much more material for a reduction of the species number.

Ranks Below Species In two ways, subspecific taxa have been created. Older authors, e.g., Linnaeus 1762, described forms deviating from the typical or α-form by adding ß or more forms indicating minor differences not demanding recognition at species level; these taxa are considered to represent varieties (ICN 2012: Article 37.4). Later authors used subspecific ranks to describe various types of differences, in most cases not applying a uniform standard. The ranks of subspecies, variety, and forma have been used extensively in those genera that are most widely grown in cultivation in Europe and America: Conophytum and Lithops. Based on definitions by Du Rietz (1930) and Stace (1985), the following guidelines are used here mainly: – The rank of subspecies is used for taxa differing only a little in their character states, growing in different geographical areas, and occurring in different ecological situations. – Varieties of a species show a lesser degree of difference than subspecies, grow rarely allopatrically but mostly in a very restricted area only, and are not necessarily ecologically differentiated. – Formae are in general sports or single aberrant plants within an otherwise rather uniform population, the difference being only expressed in a single feature, e.g., green instead of brownish leaves. In this treatment, the lowest rank considered is that of subspecies, published varieties are listed under the relevant species or subspecies.

General Scheme for the Treatment of Taxa All genera are arranged alphabetically, the name being followed by the name of the subfamily into which the genus is placed. All valid and accepted names are printed in bold, synonyms in italics. For each treatment, the contributor(s) is (are) given. The basic scheme for the treatment of an accepted taxon is as follows: – Nomenclatural part, including the name of the taxon with author and bibliographical reference(s), its synonym(s) and types with full references, etymology (only for genera) – Detailed description (see below) – Notes (see below)

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For genera, the name of the subfamily into which they are placed is stated in the headline. Each synonym appears at least twice: once together with the accepted name under which nomenclatural and bibliographical details are to be found, and secondly in the alphabetical order, either as genus or as species under the genus in which it had been placed. For subfamilial and generic synonyms a brief explanation is given for the reason(s) which render them invalid or illegitimate. The specific synonyms are principally listed together with the accepted names under the genus in alphabetical order. An exception from this rule is Mesembryanthemum for which synonyms are presented in two parts: – All names synonymous to names of taxa currently included in the genus Mesembryanthemum are given in the account itself. – All synonyms of names of taxa now placed in other genera, and those names under Mesembryanthemum that have been validly published but can at present not be placed with confidence in any of the existing other genera, are separately listed immediately after the account on the genus Mesembryanthemum.

Nomenclature and Typification Homotypic synonyms, i.e., names of taxa based on the same type, are separated by a triple hyphen (); all names are followed by the citation of their common type. A possible source of homotypic synonyms are species which have been transferred into different genera, e.g., Aspazoma amplectens (L. Bolus) N.E.Br. in Phillips 1926: 244  Mesembryanthemum amplectens L. Bolus ASAM 1913: 153 Lectotype, here designated: Pearson 6064 (BOL!). Another source are taxa that were described more than once based on the same type, as was done in Tetragonia tetragonoides and T. expansa. In contrast, heterotypic synonyms are names of taxa based on different types which are presently incorporated in a single taxon. These synonyms are separated by a double hyphen (=), and the types are cited right behind each name, e.g., Cephalophyllum parvibracteatum (L.Bolus) H.E.K.Hartmann T Frames 18967 (BOL!) = C. All valid taxa have to be typified (ICN 2012: Art. 7.1), be they legitimate or illegitimate. T here denotes a holotype, LT a lectotype, and NT a neotype. In a number of cases a lectotype or neotype is chosen here for the first time, which then is indicated as recommended (ICN 2012: Art. 7.10) by adding “here designated.” Type specimens seen are marked with an exclamation mark printed after the abbreviation of the herbarium housing the sheet, which is cited in parentheses, e.g., (BOL!).

Standard of Descriptions Descriptions are given for the family, the subfamilies, the genera, and all species and subspecies according to the state of knowledge at the date of completion of the manuscript. This implies necessarily differences in standard because for a number of genera revisions and plenty of fresh material have been

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available for this book, but numerous genera have never been revised after their establishment, though much material has been available in various herbaria. For all taxa, the original descriptions and all further combinations published have been seen (a total of about 5000 references, the few exceptions not seen are marked in the References), and results have been compared with these. In cases where revisions and plants were available, well-balanced descriptions can be offered, mostly also based on extensive field studies, which have partly been conducted for the purpose of writing the texts for this handbook. In contrast to this favorable state of knowledge, there are numerous taxa, for which no revisions were available and no studies were possible, though much material was at hand, but time was lacking. In these cases, the descriptions have been taken back to the original texts, their possibly accompanying figures, and to the types, when these have been preserved. The descriptions thus derived will disagree with many traditional usages of names, e.g., in Lampranthus, but the chosen procedure is the only basis on which to promote an unbiased study of the group. All later published descriptions are to be seen as interpretations, which may or may not be correct, as has been demonstrated in a number of cases, e.g., Lampranthus scaber (Liede & Meve 1990). In the case of complete data, the descriptions follow a scheme from growth form over leaves (L) to flowers (Fl), fruits (Fr), seeds (S), chromosomes (Chr), ecology (Ecol), and distribution (Distr). Each item is introduced by the bold letters given here in brackets. In addition, internodes (I) and inflorescences (Inf) can occur. In cases of incomplete data sets, the key words for which data are missing are left out, excepting Ecol and Distr, where “unknown” or “uncertain,” or other available information can be inserted, e.g., the continent of origin. Additional information on the taxonomic history, similarities, cultivation, and usage is given under “Note.”

Orthography Orthography is in British English; some expressions used for South African types of vegetation, e.g., “veld,” are included as well. The word “Karoo” is spelt in different ways in English. In general, the spelling “Karroo” is used in the text here; in various names, however, the original spelling like “karooides” or “karroid” is retained.

Keys Keys to the subfamilies, to their main groups, and down to the genera are given at the beginning, together with the relevant descriptions. Keys to subgenera, sections, species, and subspecies are found under the genera in those cases in which enough material has been available to construct a key. All keys are based on features that are visible through most of the year, which excludes flowers in many cases because they are short-lived and only distinctive in some taxa or when much material can be compared, which is very rarely the case.

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Distribution A map is provided for each genus, delineating the area in which material of the genus has been found, based on literature, herbarium sheets, and on well over 15,000 collections by the contributors and their collaborators. Place names are abbreviated on these maps; the complete place names referred to in distribution maps can be found on a map covering most of southern Africa (Fig. 2), which also shows the old districts of Namibia and the boundaries of the new provinces of the Republic of South Africa. In addition, for each species and subspecies the distribution is cited in the sequence of district, province, country for most parts of the world. If more than five districts in a province are included, a general description of the area is given. During the production of this work, the names of numerous districts were changed, sometimes also the boundaries. Reference is here made to the old districts by name and in their boundaries because these old names have been used in most cases in the original descriptions.

Illustrations Among the 700 figures are about 150 maps and about 550 colored images. The majority of the photographs has been taken in habitat, and it has been ascertained that the plants have been reliably identified by the authors. This procedure has the consequence that only revised or well-known taxa are figured. Since the aim was also to present plants in the field, it means that photographs taken in the open, often under difficult or adverse conditions, may not be as brilliant as those taken in a collection or garden. Species known only from literature or from herbarium material and not reconfirmed on fresh material are only described. In this way, it is hoped to break the sequence of misidentifications in several cases and at the same time to stimulate new studies in little known yet sometimes widely occurring genera like Lampranthus, Delosperma, and Ruschia. Biozentrum Klein Flottbek Hamburg, Germany

H. E. K. Hartmann

References Bittrich V. (1987) Untersuchungen zu Merkmalsbestand, Gliederung und Abgrenzung der Unterfamilie Mesembryanthemoideae (Mesembryanthemaceae Fenzl.) MIBH 21: 5–116 Bittrich V. (1990) Systematic Studies in Aizoaceae. MIBH 23b: 491–507 Bittrich V., Hartmann H.E.K. (1988) The Aizoaceae – a new approach. Botanical Journal of the Linnean Society 97: 239–254 Brown N.E. (1925) Mesembryanthemum … GC 78: 212 Chesselet P. Hartmann H.E.K. (1995) Khadia alticola Chess. & H.E.K. Hartm. spec. nov (Mesembryanthema, Aizoaceae). Aloe 32: 46–49

l Chesselet, P., Smith, G.D., van Wyk, A.E. (2001) A new tribal classification for the Mesembryanthemaceae Fenzl based on characters of the floral nectary. Aloe 38: 25–28 Chesselet, P., Smith, G.D., van Wyk, A.E. (2002) A new tribal classification of Mesembryanthemaceae: evidence from floral nectaries. Taxon 51: 295–308 Chesselet, P., Smith, G.F., van Wyk, A.E. (2005) Mesembryanthemaceae. A new tribe and adjustments to infrafamilial classification. Bothalia 34: 47–51 Dehn M. (1989) Untersuchungen zum Merkmalsbestand und zur Stellung der Gattung Antimima. N.E.Br. emend. Dehn (Mesembryanthemaceae Fenzl). MIBH 22: 189–215 Dillenius J.J. (1732) Hortus elthamensis … The author, London Dinter K. (1923a) Beiträge zur Flora von Südwestafrika I. FR 19: 122–160 (30.6.1923) Dinter K. (1923b) Beiträge zur Flora von Südwestafrika II. FR 19: 177–186 (30.7.1923) Dinter K. (1923c) Index der aus Deutsch-Südwestafrika bis zum Jahre 1917 bekannt gewordenen Pflanzenarten XI. FR 19: 187–192 (30.7.1923) Dinter K. (1923d) Succulentenforschung in Südwestafrika. I. FR Beiheft 23: 1–80 (15.8.1923) Dinter K. (1923e) Index der aus Deutsch-Südwestafrika bis zum Jahr 1917 bekannt gewordenen Pflanzenarten XII. FR 19: 235–236 (30.9.1923) Dinter K. (1928a) Südwestafrikanische Lithopsarten. R. Graessner, Perleberg Dinter K. (1928b) Sukkulentenforschung in Südwestafrika II: 12, 93, 98, 102–103, 105– 106. Gustav Winter, Herrnhuth i.Sa. (this print is also bound as FR Beiheft 53: 1–145) du Rietz G. E. (1930) The fundamental units of biolkogical taxonomy. Svensk botanisk tidskrift 24, 3 Hartmann H.E.K (1978) Monographie der Gattung Argyroderma. N.E.Br. (Mesembranthemaceae Fenzl). MIBH 15: 121–235 Hartmann H.E.K. (1983) Untersuchungen zum Merkmalsbestand und zur Taxonomie der Subtribus Leipoldtiinae (Mesembryanthemaceae). BB 136: 1–67 Hartmann H.E.K. (1988) Fruit types in Mesembryanthema. BBP 63: 313–349 Hartmann, H.E.K. (1991) Mesembryanthema. CBH 13: 75–157 Hartmann H.E.K. (1992) Ihlenfeldtia, a new genus in Mesembryanthema (Aizoaceae). BJS 114: 29–50 Hartmann H.E.K (1998a) New combinations in Ruschiodeae, based on studies in Ruschia (Aizoaceae). Bradleya 16: 44–91 Hartmann H.E.K. (1998b) New combinations in Antimima N.E.Br (Ruschiodeae, Aizoaceae). Bothalia 28: 67–82 Hartmann, H.E.K. (2001) Illustrated Handbook of Succulent Plants: Aizoaceae A-E and F-Z. Springer. Heidelberg Hartmann H.E.K (2007a) Aizoaceae in den Vereinigten Arabischen Emiraten. Avonia 25: 36–42 Hartmann H.E.K (2007b) Neues von den Aizoaceae in den Vereinigten Arabischen Emiraten. Avonia 25: 86–95 Hartmann H.E.K. (2009) A revision of the species of Delosperma (Aizoaceae) in Gauteng, South Africa. Haseltonia 15: 53–68 Hartmann, H.E.K, Niesler I.M. (2009) On the evolution of nectaries in Aizoaceae. Bradleya 27: 69–120 Haworth A.H. (1795) Observations on the genus Mesembryanthemum …, part 2. The author, London Haworth A.H. (1803) Miscellanea naturalia, … J. Taylor, London Haworth A.H. (1804) Mesembryanthemum inclaudens. Andrews Botanical Repertorium 6: t. 388 Haworth A.H. (1812) Synopsis plantarum succulentarum. R. Taylor, London Haworth A.H. (1819) Supplementum plantarum succulentarum. R. & A. Taylor. London Haworth A.H. (1821) Revisiones plantarum succulentarum. R. & A. Taylor, London Haworth A.H. (1823) Plantae Rarae Succulentae. Mesembryanthemum. Linn. &c. The Philosophical Magazine and Journal 62: 381 Haworth A.H. (1824a) Observations on the Mesembryanthema barbata. The Philosophical Magazine and Journal 64: 61–62 Haworth A.H. (1824b) An account of the Mesembryanthema Ringentia. The Philosophical Magazine and Journal 64: 109–111

Introduction

Introduction

li Haworth A.H. (1824c) Decas tertia novarum Plantarum Succulentarum. The Philosophical Magazine and Journal 64: 423–428 Haworth A.H. (1826) Decas sexta novarum Plantarum Succulentarum. The Philosophical Magazine and Journal 68: 125–132. Decas septima …: 328–331 Haworth A.H. (1827) Decas octava Plantarum Novarum Succulentarum. The Philosophical Magazine and Journal, New series 1: 124–125 Haworth A.H. (1831) Mr. Haworth's thirteenth decade of new succulent plants. The Philosophical Magazine and Journal, New series 10: 414–424 Klak C. (2010) Phylogeny and diversification of Aizoaceae: progress amd prospect. Schumannia 6, Biodiversity & Ecology 3: 87–107 Klak C., Bruyns P.V. (2013) A new infrageneric classification for Mesembryanthemum (Aizoaceae: Mesembryanthemoideae). Bothalia 43: 197–206 Klak C. et al. (2003) A phylogentic hypothesis for the Aizoaceae (Caryophllales) based on four plastid DNA regions. American Journals of Botany 90: 1433–1445 Liede S., Meve U. (1990) What is Lampranthus scaber? Bradleya 8: 39–44 Linnaeus C. (1753) Species plantarum. Facs. 1957. Ray Society, London Linnaeus C. (1762) Species plantarum, ed. 2: 515, 687–700. Salvius, Stockholm Marloth R. (1910) Some new South African succulents: Aizoaceae. TRSSA 1: 405–407, t. 27; 2,1: 33–40, t. I Marloth R. (1914) A new mimikry plant. TRSSA 4: 137–138 Niederle J. (2012a) Malotigena. Skalničkářův rok 65: 35–36 Niederle J. (2012b) Malotigena: Skalničkářův rok 66: 35 Phillips E.P. (1926): The genera of South African plants: 241–249. Cape Times, Cape Town Rowley G.D, (1978) Reunion of some genera of Mesembryanthemaceae. NCSJ 33: 6–9 – A postscript: 62 Schwantes G. (1947) System der Mesembryanthemaceen. Sukkulentenkunde 1: 34–40 Schwantes G. (1952) Die Früchte des Mesembryanthemaceen. MBMUZ 193: Series Botanik Nr. 39: 1–38 Stace, C.A. 1985, (reprint of edition from 1980 with additions), Plant Taxonomy and Biosystematics, The Bath Press, Avon Thiede J. (2004) Phylogenetics, systematics and classification of the Aizoaceae: a reconsideration based on molecular data. Schumannia 4, Biodiversity & Ecology 2: 51–58 Thiede J. (2014, submitted) Gibbaeum ihlenfeldtii (Aizoaceae), an overlooked new species from the Overberg region. Phytotaxa Thiede J., Schmidt S., Rudolph B. (2007) Phylogenetic implications of the chloroplast rpoC1 intron loss in the Aizoaceae (Caryophyllales). Biochemical Systematics and Ecology 35: 372–380

Unfortunately, the Editor-in-Chief of this volume, Heidrun Hartmann, deceased before she was able to complete the introduction to this edition. We (Wilfried Hartmann and Springer) therefore added missing references to the best of our knowledge for her. Please excuse any incorrect citations which may have crept in.

Part I The Family Aizoaceae

Aizoaceae H. E. K. Hartmann and M. Gerbaulet

Aizoaceae Martynov 1820: 15 nomen conservandum (McNeill et al. 2012; Wiersema et al. 2015: App. IIB); Bittrich & Hartmann 1988: 239–254; Bittrich 1990: 491–507; Hartmann 1991: 75–157; Hartmann 1993: 37–69; Hartmann 2001a: 1–22; Hartmann 2001b: 1–22; Klak et al. 2003: 1433–1445; Thiede 2004: 51–58 Typus Aizoon L. = Ficoidaceae Jussieu 1789: 315 nomen rejiciendum Typus Mesembryanthemum L. (as Ficoides Miller 1754: without page)
Mesembryanthemaceae Philibert 1801: 268 nomen conservandum (McNeill et al. 2012; Wiersema et al. 2015: App. IIB) Typus Mesembryanthemum L. nomen conservandum = Sesuviaceae Horaninow 1834: 83 Typus Sesuvium L. = Tetragoniaceae Lindley 1836: 209 nomen conservandum (McNeill et al. 2012; Wiersema et al. 2015: App. IIB) Typus Tetragonia L. Predominantly perennial shrubs, rarely annual to biennial herbs, rarely trees, and rarely plants reduced to a single leaf-pair, growing sometimes sunken in the ground; L mostly opposite, simple and entire, rarely lyrate, epidermis with bladder cells of various shapes and sometimes reduced, or

H.E.K. Hartmann (*) Biozentrum Klein Flottbek, Hamburg, Germany e-mail: [email protected] M. Gerbaulet Hitzacker, Germany e-mail: [email protected]

uniform and xeromorphic, often papillate to pubescent, base of petioles rarely with stipuliform appendages or leaves sessile, sometimes with a connate leaf sheath; Inf of principally dichasial pattern, complete or in various derived forms, mostly terminal, often seemingly axillary, often reduced to a single flower; Fl actinomorphic, mostly bisexual, perigynous to hypogynous or epigynous, perianth elements (3–)5(–8), free portions often unequal and with dorsal subapical appendages, basally connate and adnate to the filaments, thus forming a perianth-stamen tube, inner surface of upper portion petaloid or green, named calyx when petaloid elements are present; androecial elements 4-many, if many, the outer primordia developing often into petaloid organs (=petals), filaments rarely connate, rarely connate with the petals forming a tube, anthers dehiscing by longitudinal slits; ovary syncarpous, (1–)5 (–1)carpellate, placentation axile, basal or parietal, ovules (1–)1 per carpel, anacampylotropous or campylotropous, bitegmic, crassinucellate, rarely pendulous; Fr mostly a hygrochastic loculicidal, rarely septicidal or xeromorphic capsule, rarely schizocarpous, sometimes a hard and indehiscent 1-seeded nut, rarely a drupe, occasionally in aggregates, or a circumscissile capsule; S mostly more or less ovoid, rarely arillate, with a curved peripheral embryo, and voluminous starchy perisperm, endosperm reduced to a layer enveloping the radicle; Ecol mainly in subtropical and mediterranean climates with moderate to low

# Springer-Verlag GmbH Germany 2017 H.E.K. Hartmann (ed.), Illustrated Handbook of Succulent Plants: Aizoaceae, DOI 10.1007/978-3-662-49260-4_1

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rainfall regimes mainly as winter rains; Distr worldwide in the subtropics, the highest number of genera and the centre of diversity at species level in southwestern Africa. Note: The family name Aizoaceae was first validly published by Martynov (1820: 15) who gave a reference to an invalidly but effectively published name by Augier (1801: 129, 137–138). This supercedes the name Aizoaceae Rudolphi (1830: 53) that had for several decades been considered the correct name. In Appendix IIB of the Melbourne Code (McNeill et al. 2012; Wiersema et al. 2015), the already conserved family name Aizoaceae Martynov (against Ficoidaceae Jussieu 1789: 315) is “superconserved” against the name Mesembryanthemaceae Philibert (1801). The present treatment follows the delimitation of the family by Bittrich & Hartmann (1988) who excluded members of the family Molluginaceae. According to Klak et al. (2003: 1433–1445) and Thiede (2004: 51–58) four subfamilies can be distinguished: Aizooideae, Mesembryanthemoideae, Ruschioideae, and Sesuvioideae. The Tetragonioideae which were treated as another subfamily by Bittrich & Hartmann (1988) are in fact polyphyletic (Klak et al. 2003; Thiede 2004) with Tetragonia nested in the Aizooideae and Tribulocarpus as sistergroup to the Sesuvioideae. Tetragonia and Tribulocarpus are accordingly included in the respective subfamilies. The main synapomorphy of the family is the possession of bladder cells (Bittrich 1990: 491–507). The authors of all subfamily names are given as in Reveal (2007). 239 genera have been described as members of this family, of which 146 are accepted here.

Shrubs of many genera form the dominant part of the vegetation of the winter rainfall area of Southern Africa (e.g. Amphibolia, Aridaria, Antimima, Eberlanzia, Lampranthus, Leipoldtia, Ruschia, Vanzijlia). Some genera are also important elements of the Karroo vegetation outside this region (e.g. Ruschia, Trichodiadema), thus contributing considerably to the carrying capacity of the natural vegetation. Few species possess chemical compounds which are used in traditional healing (e.g. Sceletium, Zaleya). A selection of small-growing species (e.g. Conophytum, Lithops) is kept in horticulture, mainly indoors or in greenhouses worldwide in moderate climates. Some genera are found as garden elements or sand-binders in suitable climates (e.g. Carpobrotus, Malephora). The sour-tasting fruits of some species of Carpobrotus are sold semi-dried in the market to be used for jellies or preserves. Only one species, Tetragonia tetragonoides, is grown as a vegetable in appropriate conditions. Ficoidaceae = Aizoaceae De Jussieu (1789: 315) was the first to describe the family Ficoideae at that rank, including the genera “Reaumuria Hasselq., Nitraria Schob., Sesuvium L., Aizoon L. Ficoidea Niss., Glinus L., Orygia Forsk., Mesembryanthemum L. Ficoides T., and Tetragonia L.”. It must be noted that Ficoidea Nissole (1730: 319) is an obligate synonym of Aizoon L., whereas Ficoides Tournefort (1730: 238–241) is an obligate synonym of Mesembryanthemum L.; both names were published before Linnaeus (1753), only

Key to the Subfamilies of Aizoaceae 1. – 2. – 3. –

Petals of staminodial origin present Petals absent; perigone-lobes petaloid inside, sepaloid outside Placentation central; nectaries shell-shaped to tubular (koilomorphic); petals and stamens often united into a tube Placentation basal or parietal, extremely rarely central; nectaries crest-shaped (lophomorphic) or flat, petals mostly free Fruit a circumscissile capsule, or a woody nut which is either 5-winged or beset with numerous spines, perianth segments with subapical appendages Fruit a loculicidal or septicidal capsule, often hygrochastic, or a woody nut with ornaments principally in 4 rows, perianth segments without subapical appendages

2 3 Mesembryanthemoideae Ruschioideae Sesuvioideae Aizooideae

Aizoaceae

one was validated later: Ficoides Miller 1754. Only this name can be considered when a type species of Ficoidaceae has to be found. The question of the family name has been discussed extensively (e.g. Sprague 1922: 71; Rowley 1951: 27–28; Friedrich 1955: 56–60; Schwantes KuaS 1957: 157–158, 167–169). It was temporarily settled when Aizoaceae Rudolphi became the nomen conservandum for the relevant genera including Aizoon. The new nomen conservandum Aizoaceae Martynov does not interfere with the rejection of Ficoidaceae de Jussieu. If Mesembryanthemaceae are considered as a separate family, the name Ficoidaceae becomes an obligate synonym, since both names are based on the same genus. Mesembryanthemaceae = Aizoaceae In Appendix IIB of the Vienna Code (McNeill et al. 2006), the author of the conserved family name Mesembryanthemaceae is given as Philibert (1801: 268). For a long time, Fenzl (1836: 347, 349) had been considered the author and Mesembryanthemaceae Fenzl became a younger synonym of Aizoaceae Martynov when combined with the latter. Mesembryanthemaceae Philibert (1801), however, predates Aizoaceae Martynov (1820). This situation was dealt with by a “superconservation” of Aizoaceae Martynov in the Melbourne Code (McNeill et al. 2012; Wiersema et al. 2015). The two subfamilies Mesembryanthemoideae and Ruschioideae have often been considered as a separate family (i.e. Herre 1971) since they share a number of characters and together represent a monophylum (Bittrich 1990; Hartmann 1991; Klak et al. 2003; Thiede 2004). In that case, the family name is to be Mesembryanthemaceae Philibert. To stress the evolutionary proximity of both taxa in the context of the family Aizoaceae, Hartmann (1991: 75–157) introduced the unranked term “Mesembryanthema” for the united subfamilies Mesembryanthemoideae and Ruschioideae. Sesuviaceae = Aizoaceae Horaninow (1834: 83) was the first to use the family name “Sesuviaceae (Ficoideae)”, including the genera “Mesembryanthemum, Tetragonia, Aizoon, Sesuvium, Trianthema, Glinus;

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– Reaumuria? Nitraria?”. A little later (Horaninow 1843: 29), four tribes were listed: Paronychieae, Portulaceae, Ficoideae, and “Neuradeae et Surianeae?” containing 18 subgroups plus two with a question mark, but no genus names. Retaining the subdivision of the family as well as its circumscription more or less, Horaninow used the family name Portulacaceae for the broadened assembly of genera some years later (1847: 138–140) without giving an author or a reason for this change. The family name Sesuviaceae has very rarely been used, probably due to the fact that the family names Ficoidaceae and Aizoaceae were also available for most of the assembly of species listed by Horaninow at that time. Nevertheless, various authors refer to the “Sesuvium-group”, comprising the genera Cypselea, Sesuvium, Trianthema, and Zaleya, when investigating Aizoaceae s.lat., i.e. including Molluginaceae (e.g. Hofmann 1973). Bittrich & Hartmann (1988: 240–245) argued that the possession of bladder cells in the epidermis and the perianthstamen tube in the Sesuvium-group sensu Hofmann (1973) justify its inclusion in the family Aizoaceae as a separate subfamily Sesuvioideae. This position was confirmed by molecular studies (Klak et al. 2003; Thiede 2004). Tetragoniaceae = Aizoaceae The authorship of the family name Tetragoniaceae has been a question of some debate recently: For some time, Nakai (1942: 103) had been used as the author of the conserved family name who in turn had ascribed the name to Reichenbach in Mössler (1827: 52). In the St Louis Code (Greuter et al. 2000), Tetragoniaceae Link (1831: 17) was given. Although Link consistently used the term “order” for these names, they were traditionally treated as denoting the rank of family. However, Link (1831) also published two family names under the order Fungi, which means that the names ranked as orders throughout the work must be treated as the names of orders as well (Art. 35.5). Therefore, the name was changed again to Tetragoniaceae Lindley (1836: 209) in the Vienna Code (McNeill et al. 2006).

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If Tetragoniaceae and Aizoaceae are combined, Tetragoniaceae becomes a younger synonym of Aizoaceae.

Aizooideae H. E. K. Hartmann and M. Gerbaulet Aizooideae Sprengel ex Arnott 1832: 112; Bittrich & Hartmann 1988: 239–254; Bittrich 1990: 491–507; Hartmann 2001a: 11–12, 21; Hartmann 2001b: 11–12, 21; Klak et al. 2003: 1433–1445; Thiede 2004: 51–58 Typus Aizoon L. = Tetragonioideae Lindley 1846: 527 Typus Tetragonia L. Annual to perennial, prostrate to erect herbs, subshrubs, or shrubs, herbaceous to woody; L opposite to alternate, flat to cylindrical, mostly free, bladder hairs present; Inf mostly frondose often representing the entire plant, Fl solitary or in groups, sometimes umbellate, in cymose panicles, or in cymes, perigone 4–5-lobed, basally mostly connate, sepaloid outside, petaloid inside, nectary coating the perianth-stamen tube, stamens mostly 8 Flat base of bottom of fruit widened, containing seeds, formed like a ring-shaped pocket, septa remaining connate with valves Fruits with separate seed pockets at half the height of the capsule, septa free from valves Leaves flat, plants annual Leaves thickened, plants perennial Bladder cells reduced, only present on leaf margins and receptacle, sometimes hair-like Bladder cells not reduced, present on al green parts Locules open, seeds visible, seed pockets at outer base of locules, pedicels glabrous, leaf margins with bladder cells Locules covered by septa arching over them, seeds hidden, placenta raised, but without seed pockets, pedicel and calyx with long hairs, leaf margins with long hairs Leaves lyrate Leaves entire Expanding sheets and keels present Expanding sheets only Stigmas persisting on the capsule, woody Stigmas absent in ripe capsule Capsule opening but once, rarely again, but never repeatedly, expanding keels breaking after first opening Capsule opening repeatedly, expanding keels remaining functional Locules 5–6, valves free to base, valve wings broad, hard, erect Locules (4–6)–12, valves united laterally at base, valve wings absent Covering membranes present Covering membranes absent, sometimes narrow rims present Fruits developing on a twisting thin stalk, with a distinct breaking-point below the base, breaking off when ripe, tumble fruits more or less globose Fruit development different Covering membranes of constant shape, even after the seeds have been removed Covering membranes declining into empty locules Expanding keels merging into expanding sheets Expanding keels distinctly separated from expanding sheets, keels parallel, high, more or less sharply pleated

2 3 Carpobrotus Sarcozona 4 6 5 Hymenogyne Caryotophora Ruschianthemum 7 9 8 Conicosia Saphesia Skiatophytum 10 15 11 12 Apatesia Carpanthea Aethephyllum 13 Cleretum 14 Dorotheanthus Pherolobus 16 17 Stoeberia Stayneria 18 20 Brianhuntleya 19 48 49 21 22 (continued)

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21. – 22. – 23. – 24. – 25. – 26. – 27. – 28. – 29. – 30. – 31. – 32. – 33. – 34. – 35. – 36. – 37. – 38. – 39. – 40. – 41. – 42. –

H.E.K. Hartmann et al.

Heterophyllous shrubs Homophyllous plants Calyx and leaves sticky, sand adhering to it Calyx and leaves not sticky Valve wings absent or as awns Valve wings present, more or less rectangular Capsule opening incompletely (valves at most erect), breaking into mericarps early, leaves more or less club-shaped with apical windows Capsule opening completely (valves spreading) and persistent, leaves without apical windows Pedicel very long (5–12 cm), flowers mostly ternate, leaves keeled, toothed, plants compact Pedicel shorter (to 2.5 cm), flowers solitary, leaves not keeled, not toothed, plants shrubby, caespitose to decumbent or creeping Shrubs, flowers opening midday Plants creeping or caespitose, flowers opening at night Valve wings at most as broad as expanding keels Valve wings broader than expanding keels Plants creeping Plants compact to caespitose Bracteoles present (different from foliage leaves) Bracteoles absent (indistinguishable from foliage leaves) Base of capsule with very long soft hairs, leaves hairy Base of capsule and leaves not velvety but glabrous, with bladder cells, or papillate Flowers with a short hypanthium Flowers with free elements from above the ovary Leaves smooth, glabrous Leaves finely granulate and rough Plants mat-forming, if more compact, leaves trigonous in section Plants compact, leaves mostly dentate, if not, leaves apically broadly widened Base of capsule and leaves with bladder cells Base of capsule and leaves xeromorphic, glabrous Plants heterophyllous, always compact Plants homophyllous Plants caespitose to shrubby, locules 5 Plants forming dense mats, adventitious roots present, locules 5–8 Leaves obliquely triquetrous Leaves isometric Plants caespitose Plants compact Plants forming clumps above the ground Plants sunken in the ground Predominant number of locules per fruit in a population 5 or 6 Predominant number of locules per fruit 7 or more Seeds tuberculate, tubercles distant Seeds smooth to rugose Flowers in inflorescences raised well above the plant Flowers solitary

Mitrophyllum 46 Psammophora 23 24 27 Frithia 25 Carruanthus 26 Jensenobotrya Neohenricia 28 29 Mossia 47 42 30 Gibbaeum 31 32 34 Peersia 33 Rhinephyllum Stomatium 35 37 Oophytum 36 Delosperma Malotigena Schwantesia 38 Corpuscularia 39 Dinteranthus 40 Lithops 41 Dinteranthus Lithops 43 44 (continued)

Aizoaceae 43. – 44. – 45. – 46. – 47. – 48. –

49. – 50. – 51. – 52. – 53. – 54. – 55. – 56. – 57. – 58. – 59. – 60. – 61. – 62. – 63. – 64. –

Flowers remaining open during and after anthesis, petals stiff Flowers closing over night, petals rather lax Locules  7 Locules  10 Plants above ground, leaves almost semi-globose, more or less appressed Plants at least partly sunken in the ground, leaves triangular and pointed, spreading Locules mostly 5, stamens > 10 Locules 4, stamens < 10 Capsules dark brown, top rounded Capsules light brown, top flat Covering membranes without distal recurving and always without any additional closing devices like rodlets, ledges or bulges at their distal ends Covering membranes with a more or less distinct distal recurving, with or without additional closing devices like rodlets, ledges or bulges, if straight, always with some additional closing device Covering membranes declining into the empty locules along a preformed edge Covering membranes straight or only slightly and gradually declining into empty locules Plants with distinct prominent bladder cells, glistening in the sun, often appearing scaly on herbarium material Plants rough, glabrous or hairy Plants heterophyllous Plants homophyllous Expanding keels divergent, emerging gradually from expanding sheets Expanding keels parallel, sheets separate from expanding keels Plants compact and low, < 4 cm h Plants shrubby, if compact, > 5 cm h Basal parts of old leaf sheaths sclerotic, completely enveloping the stem Old leaf-sheaths papery-pergamentaceous Flowers basally enclosed by connate hypsophylls forming a cup Flowers exserted above leaves, no cups Both leaf pairs of a season connate for > 30% One leaf pair of a season connate for < 25%, spreading, the other for > 50%, erect Leaves with an apical diadem Leaves without an apical diadem Bladder cells on leaves thin-walled, collapsing when dry, surface glittering Bladder cells thick-walled, constant in shape, surface dull Surfaces of primary internodes with prominent bladder cells Surfaces of primary internodes glabrous Leaves fat-digitiform, > 4 cm l Leaves subterete to trigonous, < 3 cm l Locules 5 Locules 6–11 Locules  8 Locules 6 Flowers white over pink to purple Flowers yellow Fruits brown, hard Fruits whitish to light brown, more or less soft

15 Nelia Hartmanthus Conophytum 45 Namibia Nananthus Delosperma Ectotropis Antegibbaeum Ruschianthus 49 91

78 50 51 63 52 57 53 Oophytum Diplosoma 54 Monilaria 55 Dicrocaulon 56 Meyerophytum Mitrophyllum Trichodiadema 58 59 62 Drosanthemum 60 Jacobsenia 61 Mestoklema Malephora Jacobsenia Knersia 64 72 65 69 (continued)

16 65. – 66. – 67. – 68. – 69. – 70. – 71. – 72. – 73. – 74. – 75. – 76. – 77. – 78. – 79. – 80. – 81. – 82. – 83. – 84. – 85. – 86. – 87. –

H.E.K. Hartmann et al. Fruits in raised inflorescences Fruits solitary Plants shrubby Plants compact to caespitose Leaves digitiform, more than 3x as long as broad, valve wings broad Leaves more or less trigonous, with convex sides, plants compact, valve wings narrow to absent Plants with thick tap-roots Plants with thinner roots Erect shrubs Compact, creeping, or caespitose plants Calyx lobes 4 Calyx lobes 5 Flowers bracteolate Flowers ebracteolate Closing body bipartite Closing body absent or present as a single body Closing body a broad sill Closing body absent or present as a round body Fruits on short stalks, closing body big and round Fruits on long stalks, closing body absent or small At least some leaves with teeth along keel and/or margins All leaves without teeth Leaves soft, fruits whitish, locules always with false septa Leaves firm, fruits brown, locules without false septa Nectary a crenulate ring Nectary as 6 glands Valve wings absent, rims of valves high Valve wings present, at least basally very broad Capsules brown Capsules whitish-grey Fruits persisting on rather long stalks Fruits breaking off their stalks, dispersed as tumble fruits Leaves velvety Leaves smooth to rough from warts Leaves sharply pointed and triangular in section, surface finely warty Leaves flattened or rounded at tips and margins, surface with prominent warts Closing body present Closing body absent Base of capsule papillate, leaves dotted Base of capsule smooth, leaves smooth Base of capsule papillate; leaves papillate, keeled, toothed Base of capsule smooth, leaves smooth or warty, never toothed Locules 5–7, rarely in single fruits 8 Locules  8 in all fruits Locules very shallow, covering membranes therefore held up by the funicles; flowers appearing after the leaf pairs thus seemingly lateral Locules deeper, covering membranes bending into empty locule; flowers distinctly terminal

Nelia 66 Polymita 67 Antegibbaeum 68 Ebracteola 90 70 71 Juttadinteria Hartmanthus Hammeria 77 Rhombophyllum 73 Bergeranthus 74 Bijlia 75 Chasmatophyllum 76 Malephora Hereroa Disphyma Gibbaeum Ebracteola 79 80 83 Prepodesma 81 Deilanthe 82 Rabiea Aloinopsis 84 85 Ihlenfeldtia Tanquana Vanheerdea 86 87 89 Didymaotus 88 (continued)

Aizoaceae 88. – 89. – 90. – 91. – 92. – 93. – 94. – 95. – 96. – 97. – 98. – 99. – 100. – 101. – 102. – 103. – 104. – 105. – 106. – 107. – 108. – 109. –

Leaves warty, plants mostly sunken in the ground Leaves deltoid, keeled, gibbose, plants above ground Shrubs Compact plants Top of capsule convex from high valve rims Top of capsule flat Closing body absent Closing body present, sometimes as a broad sill Valve wings, at least at base, much broader than expanding keels Valve wings absent or narrow, i.e.  width of expanding keels Valve wings tapering into awns towards the tip of the valve Valve wings rectangular Capsules multilocular, plants compact Capsules 5-locular, if multilocular, plants shrubby Valve wings in open capsules stiff, stretching over the locules and not moving outwards with the valves Valve wings in open capsules flexible, moving outwards with the valve Seeds echinate Seeds with low papillae Plants compact Plants shrubby, rarely caespitose or creeping Filaments broadened and connate at base Filaments filiform, free Leaves mesomorphic Leaves xeromorphic Flowers yellow Flowers white to purple Seeds echinate Seeds smooth to moderately papillate Surface of ovary concave, forming a hypanthium in the flower Surface of ovary flat to raised Leaves connate into a sheath which is continuous with the cortex of the preceding internode Leaves more or less free or only somewhat fused Free parts of leaves < 10 mm l Free parts of leaves 20–30 mm l Fruits reddish-brown, not closing again completely after the first opening, shrubs to 2mh Fruits with a white cover on top, closing completely again after the first opening, shrubs to c. 40 cm h Flowers white to pink, leaves grey from an extremely thick wax cover Flowers purple or magenta, leaves bright green Leaves club-shaped, apically rounded, flowers ebracteolate, 4 calyx lobes Leaves with a marked tooth or mucro at the tip, flowers with bracteoles, 5 calyx lobes Closing body bipartite Closing body not divided Closing body U-shaped, leaves mesomorphic, plants creeping Closing body of two more or less flat plates, leaves xeromorphic, plants compact to shrubby

17 Titanopsis Lapidaria Zeuktophyllum Dracophilus Cerochlamys Vlokia 92 108 93 99 94 95 Pleiospilos Erepsia Faucaria 96 Braunsia 97 Orthopterum 98 Circandra Lampranthus 100 101 Scopelogena Oscularia Namaquanthus 102 Erepsia 103 104 106 Smicrostigma 105 Stayneria Zeuktophyllum Wooleya 107 Enarganthe Esterhuysenia 109 110 Disphyma Rhombophyllum (continued)

18 110. – 111. – 112. – 113. – 114. – 115. – 116. – 117. – 118. – 119. – 120. – 121. – 122. – 123. – 124. – 125. – 126. – 127. – 128. – 129. – 130. – 131. – 132. – 133. –

H.E.K. Hartmann et al. Leaves soft, mesomorphic with bladder cells Leaves hard, xeromorphic without bladder cells Covering membranes more or less straight, plants heterophyllous Covering membranes undulate, leaves anisophyllous in a pair Expanding keel clearly separate from expanding sheet Expanding keel merging into expanding sheet Valve wings broad (at least at base) Valve wings absent or narrow ( width of expanding keels) Valve wings more or less rectangular Valve wings tapering from a broad base Closing bodies not larger than seeds, leaving space at the sides Closing bodies larger than seeds, more or less blocking the distal opening of the locule Capsule multilocular Capsule pentamerous Capsules dispersed as tumble fruits, leaves with apical window, plants under ground Capsules persistent on pedicels, leaves fusiform without a window, plants above ground Top of capsule with remains of hard bases of stigmata in the centre Top of capsule flat in the centre Base of capsule velvety or rough from papilla Base of capsule glabrous First leaves of side branches connate > 50% First leaves of side branches hardly connate like all others (seedlings occasionally with connate leaf pairs) Capsules with closing bulges Capsules with closing rodlets Top of capsule turret-shaped; procumbent branches from compact centre not rooting Top of capsule rounded, but not turret-shaped, erect to ascending, or saltatory and rooting 5 locules Many locules Leaves sharply triquetrous, grey Leaves trigonous to semi-ovate, gibbose, dark green Closing body large, completely locking the locule Closing body smaller, leaving some space at the sides Capsule pentamerous Capsule multilocular Flowers yellow Flowers pink to purple Leaves awl-shaped, the keel visible as a line in turgid conditions Leaves never awl-shaped, a keel always prominently visible Leaves glabrous Leaves velvety, hairy or rough Flowers with a hypanthium; capsule 12–24 locular Flowers without a hypanthium; capsule 8-locular Leaves and base of capsule velvety Leaves and base of capsule rough Flowers with c. 2000 elements Flowers with up to 300 elements Flowers yellow, shrubs compact Flowers white to purple

111 112 Mitrophyllum Glottiphyllum 113 Bijlia 114 125 115 123 116 119 117 118 Fenestraria Jordaaniella Arenifera Amphibolia Cheiridopsis 120 Vanzijlia 121 Cephalophyllum 122 Hallianthus Leipoldtia Eberlanzia 124 Machairophyllum Pleiospilos 126 132 127 129 Bergeranthus 128 Marlothistella Antimima 130 131 Argyroderma Octopoma Odontophorus Cheiridopsis Cylindrophyllum 133 134 135 (continued)

Aizoaceae 134. – 135. – 136. – 137. – 138. – 139. – 140. – 141. – 142. –

19

Leaves sharply triquetrous, grey Leaves scimitar-shaped with a rounded keel Plants compact, caespitose or creeping Shrubs Petals white or purple, never striped Petals with a central, longitudinal deep purple stripe, i.e. vittate Seeds echinate or markedly papillate Seeds smooth to rough Flowers in annually enlarged, persisting inflorescences Flowers solitary or in annually formed and ripening inflorescences, new ones formed every year Fruits mostly 5, rarely 6 locules Fruits with mostly 8, rarely 7–11 locules Valves without wings Valves with narrow wings broadest in their middle Calyx lobes 4–5, prominent closing rodlet present Calyx lobes 6, sharp closing ledge on distal rim Leaves of a pair basally nearly free Leaves of a pair basally connate, forming a short sheath

Machairophyllum Hereroa 136 137 Khadia Acrodon Astridia 138 Ottosonderia 139 140 142 Ruschia 141 Ruschiella Schlechteranthus Lampranthus Phiambolia

Key to the Genera of Sesuvioideae 1. – 2. – 3. – 4. –

Fruit a nut Fruit a circumscissile capsule Style 1 Styles 2–5 Septa complete in ripe fruits; seeds brown to blackish Septa incomplete or absent in ripe fruits; seeds light coloured Operculum of capsule breaking into 2 parts, 4 seeds per fruit Operculum of capsule in one piece; numerous seeds per fruit

part), denoting the shape of the nectary, and ultimately six (1960) subtribes (Rappa & Camarrone 1953, 1960). All names below subfamily are also invalid. Since all the genera cited under Lophomorphoideae Rappa & Camarrone 1953 and 1960 are now placed in the subfamily Ruschioideae, the former name is a synonym of the latter.

Sesuvioideae H. E. K. Hartmann and M. Gerbaulet Sesuvioideae Lindley 1846: 527; Adamson 1962: 243–253; Bittrich & Hartmann 1988: 239–254; Bittrich 1990: 491–507; Hartmann 2001a: 21; Hartmann 2001b: 21; Klak et al. 2003:

Tribulocarpus 2 Trianthema 3 4 Cypselea Zaleya Sesuvium

1433–1445; Thiede 2004: 51–58
Sesuviaceae Horaninow 1834: 83 Typus Sesuvium L. Prostrate to erect perennials or annuals; L flat to terete, more or less unequal, with or without stipule-like appendages, often with membranous sheaths at the base of the petiole, leaf surface with more or less prominent bladder cells, hairy or smooth; Inf terminal but apparently axillary through sympodial branching, Fl solitary or in clusters, rarely in somewhat loose dichasia, perianth with subapical appendages with unifacial tips, rarely without, perigynous, 1–5 styles, ovules 2-many; Fr a circumscissile capsule or a nut; S completely enveloped by an aril; Chr x = 8; Ecol coastal to inland in mostly moister climates than the other subfamilies; Distr worldwide in the tropics and subtropics.

20

Note: Based on molecular studies (Klak et al. 2003: 1433–1445; Thiede 2004: 51–58), the genus Tribulocarpus (Tetragonoidedae) is now included in the Sesuvioideae. Five genera are distinguished. Members of the subfamily are probably best known from tropical coasts in brackish soil, sometimes on rocks, often near mangrove, where the plants can form conspicuous ground covers, red to purple in many cases. Other members are found in disturbed areas near roads, being typical weeds, or in low and open deciduous bushland or woodland.

References Adamson R.S. (1962) The South African species of Aizoaceae XII. Sesuvium, Trianthema and Zaleya. JSAB 28: 243–253 Arnott G.A.W. (1832) Botany: 112. Preprint of an article on botany for the 7th ed. of the Encyclopaedia Britannica in 1842 Augier A. (1801) Essai d’une nouvelle classification des végétaux. Bruyset Ainé, Lyon Bittrich V. (1987) Untersuchungen zu Merkmalsbestand, Gliederung und Abgrenzung der Unterfamilie Mesembryanthemoideae (Mesembryanthemaceae FENZL). MIBH 21: 5–116 Bittrich V. (1990) Systematic studies in Aizoaceae. MIBH 23b: 491–507 Bittrich V., Hartmann H.E.K. (1988) The Aizoaceae – a new approach. Botanical Journal of the Linnean Society 97: 239–254 Brown N.E. (1925) Mesembryanthemum . . . GC 78: 232, 412–413, 433, 451, 468, 484, 500 Burnett G.T. (1835) Outlines of Botany. Henry Renshaw, London Candolle A.P. de (1828) Prodromus systematis naturalis regni vegetabilis, sive . . . 3: 352, 359, 415–456. Treuttel & Würtz, Paris Chesselet P., Hartmann H.E.K. (1995) Khadia alticola Chess. & H.E.K. Hartm. spec. nov. (Mesembryanthema, Aizoaceae). Aloe 32: 46–49 Chesselet, P., Smith, G.D., van Wyk, A.E. (2001) A new tribal classification for the Mesembryanthemaceae Fenzl based on characters of the floral nectary. Aloe 38: 25–28 Chesselet P., Smith G.F., van Wyk A.E. (2002) A New Tribal Classification of Mesembryanthemaceae: Evidence from Floral Nectaries. Taxon 51: 295–308 Chesselet P., van Wyk A.E., Smith G.F. (2004) Mesembryanthemaceae. A new tribe and adjustments to infrafamilial classification. Bothalia 34: 47–51

H.E.K. Hartmann et al. Dehn M. (1989) Untersuchungen zum Merkmalsbestand und zur Stellung der Gattung Antimima N.E.Br. emend. Dehn (Mesembryanthemaceae Fenzl). MIBH 22: 189–215 Dehn M. (1993) Untersuchungen zum Verwandtschaftskreis der Ruschiinae (Mesembryanthemaceae) MIBH 24: 91–198 Dillenius J.J. (1732) Hortus elthamensis . . . The author, London Dinter K. (1923a) Beiträge zur Flora von Südwestafrika I. FR 19: 122–160 (30.6.1923) Dinter K. (1923b) Beiträge zur Flora von Südwestafrika II. FR 19: 177–186 (30.7.1923) Dinter K. (1923c) Index der aus Deutsch-Südwestafrika bis zum Jahre 1917 bekannt gewordenen Pflanzenarten XI. FR 19: 187–192 (30.7.1923) Dinter K. (1923d) Succulentenforschung in Südwestafrika. I. FR Beiheft 23: 1–80 (15.8.1923) Dinter K. (1923e) Index der aus Deutsch-Südwestafrika bis zum Jahr 1917 bekannt gewordenen Pflanzenarten XII. FR 19: 235–236 (30.9.1923) Dinter K. (1928a) Südwestafrikanische Lithopsarten. R. Graessner, Perleberg Dinter K. (1928b) Sukkulentenforschung in Südwestafrika II: 12, 93, 98, 102–103, 105–106. Gustav Winter, Herrnhuth i.Sa. (this print is also bound as FR Beiheft 53: 1–145) Du Rietz G.E. (1930) The fundamental units of biological taxonomy. Svensk Botanisk Tidskrift 24: 333–428 Fenzl E. (1836) Monographie der Mollugineen und Steudelieen, zweier Unterabteilungen der Familie Portulaceen. Annalen des Wiener Museums für Naturgeschichte 1: 337–386 Friedrich H.-C. (1955) Beiträge zur Kenntnis einiger Familien der Centrospermae. MBSM 12: 56–60 Gerbaulet M. (2012) One or many genera in Mesembryanthemoideae (Aizoaceae)? Discussion of a conflict in genus perception. Bradleya 30: 187–198 Gerbaulet M., Struck M. (2001a) In: Hartmann, H.E.K. 2001a: Illustrated Handbook of Succulent Plants: Aizoaceae A-E: Aptenioideae: 12; Coilomorphioideae: 12; Mesembryanthemoideae: 13–14. Springer, Heidelberg Gerbaulet M., Struck M. (2001b) In: Hartmann H.E.K. 2001b: Illustrated Handbook of Succulent Plants: Aizoaceae F-Z: Aptenioideae: 12; Coilomorphioideae: 12; Mesembryanthemoideae: 13–14. Springer, Heidelberg Greuter W., McNeill J., Barrie F.R., Burdet H.M., Demoulin V., Figueiras T.S., Nicolson D.H., Silva J. E., Skog P.C., Trehane P., Turland N.J., Hawksworth D.L. (2000) ICBN International Code of Botanical Nomenclature (St Louis Code). Regnum Vegetabile 138, Koeltz, Königstein Hartmann H.E.K. (1978) Monographie der Gattung Argyroderma N.E.Br. (Mesembryanthemaceae Fenzl). MIBH 15: 121–235

Aizoaceae Hartmann H.E.K. (1983) Untersuchungen zum Merkmalsbestand und zur Taxonomie der Subtribus Leipoldtiinae (Mesembryanthemaceae). BB 136: 1–67 Hartmann H. (1988) Fruit types in Mesembryanthema. BBP 63: 313–349 Hartmann H.E.K. (1991) Mesembryanthema. CBH 13: 75–157 Hartmann H.E.K. (1992) Ihlenfeldtia, a new genus in Mesembryanthema (Aizoaceae). BJS 114: 29–50 Hartmann H.E.K. (1993) Aizoaceae. In: Kubitzki K.: The families and genera of vascular plants: 37–69. Springer, Berlin Heidelberg New York Hartmann H.E.K. (1998a) New combinations in Ruschioideae, based on studies in Ruschia (Aizoaceae). Bradleya 16: 44–91 Hartmann H.E.K. (1998b) New combinations in Antimima N.E.Br. (Ruschioideae, Aizoaceae). Bothalia 28: 67–82 Hartmann H.E.K. (ed.) (2001a) Illustrated Handbook of Succulent Plants: Aizoaceae A-E. Springer, Heidelberg Hartmann H.E.K. (ed.) (2001b) Illustrated Handbook of Succulent Plants: Aizoaceae F-Z. Springer, Heidelberg Hartmann H.E.K. (2007a) Aizoaceae in den Vereinigten Arabischen Emiraten. Avonia 25: 36–42 Hartmann H.E.K. (2007b) Neues von den Aizoaceae in den Vereinigten Arabischen Emiraten. Avonia 25: 86–95 Hartmann H.E.K., Bittrich V. (1990) Nomenclature in Mesembryanthema (Aizoaceae): The generic names by Rappa & Camarrone. Bothalia 20: 153–157 Hartmann H.E.K., Bittrich V. (1991) Typification of suprageneric names – some nomenclatural changes in Aizoaceae. SAJB 57: 73–74 Hartmann H.E.K., Liede-Schumann S. (2014) Two new subgenera and one new species in the genus Drosanthemum. Bradleya 32: 50–63 Hartmann, H.E.K., Niesler, I.M. (2009): On the evolution of nectaries in Aizoaceae. Bradleya 27: 69–120 Haworth A.H. (1795) Observations on the genus Mesembryanthemum . . ., part 2. The author, London Haworth A.H. (1803) Miscellanea naturalia, . . . J. Taylor, London Haworth A.H. (1804) Mesembryanthemum inclaudens. Andrews Botanical Repertorium 6: t. 388 Haworth A.H. (1812) Synopsis plantarum succulentarum. R. Taylor, London Haworth A.H. (1819) Supplementum plantarum succulentarum. R. & A. Taylor. London Haworth A.H. (1821) Revisiones plantarum succulentarum. R. & A. Taylor, London Haworth A.H. (1823) Plantae Rarae Succulentae. Mesembryanthemum. Linn. &c. The Philosophical Magazine and Journal 62: 381 Haworth A.H. (1824a) Observations on the Mesembryanthema barbata. The Philosophical Magazine and Journal 64: 61–62 Haworth A.H. (1824b) An account of the Mesembryanthema Ringentia. The Philosophical Magazine and Journal 64: 109–111

21 Haworth A.H. (1824c) Decas tertia novarum Plantarum Succulentarum. The Philosophical Magazine and Journal 64: 423–428 Haworth A.H. (1826) Decas sexta novarum Plantarum Succulentarum. The Philosophical Magazine and Journal 68: 125–132. Decas septima . . .: 328–331 Haworth A.H. (1827) Decas octava Plantarum Novarum Succulentarum. The Philosophical Magazine and Journal, New series 1: 124–125 Haworth A.H. (1831) Mr. Haworth’s thirteenth decade of new succulent plants. The Philosophical Magazine and Journal, New series 10: 414–424 Herre H. (1971) The Genera of Mesembryanthemaceae. Tafelberg, Cape Town Hofmann U. (1973) Morphologische Untersuchungen zur Umgrenzung und Gliederung der Aizoaceae. BJS 93: 247–324 Horaninow P.F. (1834) Primae Lineae Systematis Naturae, . . .: 83. Krajan, St. Petersburg Horaninow P.F. (1843) Tetractys naturae seu systema quadrimembre omnium naturalium, quod primis lineis systematis naturae, a se editis: 29. Wienhöber, St. Petersburg Horaninow P.F. (1847) Characteres essentiales familiarum ac tribuum regni vegetabilis et amphorganici ad leges tetracydis naturae conscripti . . .: Portulacaceae: 138–140. Wienhöber, St. Petersburg Ihlenfeldt H.-D., Gerbaulet M. (1990) Untersuchungen zum Merkmalsbestand und zur Taxonomie der Gattungen Apatesia N.E.Br., Carpanthea N.E.Br., Conicosia N.E.Br., Herrea Schwantes und Hymenogyne Haw. (Mesembryanthemaceae Fenzl) BJS 11: 457–498 Ihlenfeldt, H.-D., Hartmann, H. (1982) Leaf surfaces in Mesembryanthemaceae. In: Cutler, D. F., Alvin, K. L., Price, C. E. (eds.): The plant cuticle. Linnean Society Symposium Series 10. 397–424 Ihlenfeldt H.-D., Schwantes G., Straka H. (1962) Die höheren Taxa der Mesembryanthemaceae. Taxon 11: 52–56 Ihlenfeldt H.-D., Struck M. (1987) Morphologie und Taxonomie der Dorotheanthinae Schwantes (Mesembryanthemaceae). BBP 61: 411–453 Jacobsen H. (1960) A handbook of succulent plants 3. Mesembryanthemums. Blandford, London Jacobsen H. (1970) Das Sukkulentenlexikon. Fischer, Jena Jacobsen H. (1974) Lexicon of succulent plants. Blandford, London Jacobsen H. (1981) Das Sukkulentenlexikon, ed. 2. Fischer, Jena Jürgens N. (1990) A life form concept including anatomical characters, adapted for the description of succulent plants. MIBH 23a: 321–341 Jussieu A.L. de (1789) Genera Plantarum 15: 312–317. Herissont & Barrois, Paris Klak C. (2010) Phylogeny and diversification of Aizoaceae: progress and prospects. Schumannia 6, Biodiversity & Ecology 3: 87–107

22 Klak C., Bruyns P.V. (2012) Phylogeny of the Dorotheantheae (Aizoaceae), a tribe of succulent annuals. Taxon 61: 293–307 Klak C., Bruyns P.V. (2013) A new infrageneric classification for Mesembryanthemum (Aizoaceae: Mesembryanthemoideae). Bothalia 43: 197–206 Klak, C., Bruyns, P.V., Hanaček, P. (2013) A phylogenetic hypothesis for the recently diversified Ruschieae (Aizoaceae) in southern Africa. Molecular Phylogenetics and Evolution 69: 1005–1020 Klak C., Bruyns P.V., Hedderson T.A.J. (2007) A phylogeny and new classification for Mesembryanthemoideae (Aizoaceae). Taxon 56: 737–756 Klak C., Khunou A., Reeves G., Hedderson T.A.J. (2003) A phylogenetic hypothesis for the Aizoaceae (Caryophyllales) based on four plastid DNA regions. American Journal of Botany 90: 1433–1445 Liede S., Meve U. (1990) What is Lampranthus scaber? Bradleya 8: 39–44 Lindley J. (1836) A Natural System of Botany. Longman, Rees, Orme, Brown, Green, & Longman, London. Lindley J. (1846) The Vegetable Kingdom. Bradbury & Evans, London Link H.F. (1829–1833) Handbuch zur Erkennung der nutzbarsten und am häufigsten vorkommenden Gewächse 2: 12–18. Haude & Spenersche Buchhandlung (S.J. Josephy), Berlin (following Stafleu & Cowan, 1981, Taxomomic Literature II: 69, this volume came out in Jan.–Aug. 1829; following Brizicky G. (1969) In Taxon 18: 651, the volume appeared in 1831) Linnaeus C. (1753) Species plantarum. Facs. 1957. Ray Society, London Linnaeus C. (1762) Species plantarum, ed. 2: 515, 687–700. Salvius, Stockholm Marloth R. (1910) Some new South African succulents: Aizoaceae. TRSSA 1: 405–407, t. 27; 2,1: 33–40, t. I Marloth R. (1914) A new mimikry plant. TRSSA 4: 137–138 Martynov I.I. (1820) Tekhno-Botanicheskii Slovar, ia Latinskom i Rossilskom Yazuikakh: 15. Rossilska Akademia, St. Peterburgh McNeill J., Barrie F.R., Burdet H.M., Demoulin V., Hawksworth D.L. Marhold K., Nicolson D.H., Prado J., Silva P.C., Skog J.E., Wiersema J.H., Turland N.J. (2006) International Code of Botanical Nomenclature (Vienna Code). Regnum Vegetabile 146: [i]-xviii, 1–568 McNeill J., Barrie F.R., Buck W.R., Demoulin V., Greuter W., Hawksworth D.L. Herendeen P.S., Knapp S., Marhold K., Prado J., Prud’homme van Reine W.F., Smith, G.F., Wiersema J.H., Turland N.J. (2012) ICN International Code of Nomenclature for algae, fungi and plants (Melbourne Code). Regnum Vegetabile 154: 1–140 Miller P. (1754) The gardeners dictionary . . . abridged from the last folio edition, . . . 2: Ficoides, 3: Tetragonocarpos (no pages given). Rivington, London Nakai T. (1942) Notulae ad plantas Asiae Orientalis (18). Journal of Japanese Botany 18: 91–103 Niederle J. (2012a) Malotigena. Skalničkářův rok 65: 35–36

H.E.K. Hartmann et al. Niederle J. (2012b) Malotigena. Skalničkářův rok 66: 35 Nissole G. (1730) Etablissement De Quelques Nouveaux Genres De Plantes. Histoire de l’Académie Royale des Sciences of the year 1711: 316–320, 324, t. 13, Paris Philibert J.-C. (1801) Introduction à l’étude de la botanique ed. 2, 3. Digeon, Paris Rappa F., Camarrone V. (1953) La nova sistematica di Mesembrianthemum. Lavori dell’Istituto Botanico e del Giardino Coloniale di Palermo 14: 1–39 Rappa F., Camarrone V. (1955) Divinazioni mendelejeffiane in sistematica botanica. Lavori dell’Istituto Botanico e del Giardino Coloniale di Palermo 15: 1–16 Rappa F., Camarrone V. (1960) La Classificazione naturale delle Mesembriantemacee. Lavori dell’Istituto Botanico e del Giardino Coloniale di Palermo 18: 11–32; also as reprint available: 1–24 Reichenbach H.G.L. (1827) Mösslers Handbuch der Gewächskunde . . . ed. 2, 1: 51–52. Hammerich, Altona Reveal J.L. (2007) Indices Nominum Supragenericorum Plantarum Vascularium. Online Rowley G.D. (1951) Studies in Ficoidaceae. I. The family name. NCSJ 6: 27–28 Rowley G.D. (1978) Reunion of some genera of Mesembryanthemaceae. NCSJ 33: 6–9; – A postscript: 62 Rudolphi F. (1830) Systema orbis vegetabilium. Gryphiae. Schwantes G. (1947) System der Mesembryanthemaceen. SKK 1: 34–40; also as reprint available: 1–7 Schwantes G. (1952) Die Früchte der Mesembryanthemaceen. MBMUZ 193: Serie Botanik Nr. 39: 1–38 Schwantes G. (1957) Ficoidaceae (Juss.) em. Hutchinson. KuaS 8: 157–158, 167–169 Schwantes G. (1960) In: Jacobsen H. A handbook of succulent plants 3. Mesembryanthemums. Blandford, London Sprague T.A. (1922) The nomenclature of plant families. JB 60: 69–73 Stace C.A. (1985) Plant Taxonomy and Biosystematics. Edward Arnold, London. Straka H. (1955) Anatomische und entwicklungsgeschichtliche Untersuchungen an Früchten paraspermer Mesembryanthemen. Nova Acta Leopoldina Neue Folge 118: 127–190 Thiede J. (2004) Phylogenetics, systematics and classification of the Aizoaceae: a reconsideration based on molecular data. Schumannia 4/Biodiversity & Ecology 2: 51–58 Thiede J., Schmidt S., Rudolph B. (2007) Phylogenetic implication of the chloroplast rpoC1 intron loss in the Aizoaceae (Caryophyllales). Biochemical Systematics and Ecology 35: 372–380 Tournefort J.P. de (1730) Etablissement de quelques nouveaux genres de plantes. Histoire de l’Académie Royale des Science 1705: 236–241, t. 4 Wiersema J.E.H., McNeill J., Turland N., Barrie F.R., Buck W.R., Demoulin V., Greuter W., Hawksworth D., Herendeen P., Knapp S., Marhold K., Prado J., Prud’homme van Reine W.F., Smith G.F. (2015) International Code of Nomenclature for algae, fungi, and plants (Melbourne Code) – Appendices II-VIII. Regnum Vegetabile 157: 429 pp.

Part II Genera of the Family Aizoaceae

Acrodon RUSCHIOIDEAE H. E. K. Hartmann

Acrodon N.E.Br. GC 1927: 12; Glen 1986: 203–226; Hartmann 1996: 29–56; Hartmann 2001: 23–25; Hartmann & Niesler 2012: 33–60 Typus A. bellidiflorus (L.) N.E.Br. Etym Gk akros, point, tip, apex odons, tooth, referring to the acute, apically dentate leaves. Compact, caespitose, or procumbent; L basally connate forming a short persisting sheath,, triquetrous with dentate keel and often dentate margins, 13–100 mm l, teeth big and flexible, epidermis cells flat to somewhat raised with thick outer walls filled with crystal sand, wax in thin horizontal plates broken into smaller ones resulting in a rugose surface; Fl 1–3, mostly with basal bracteoles, pedicels 15–40 mm l, K 5, almost equal, petals white, rarely pink, apically purple with a central, longitudinal red-purple stripe (vittate), often also with a purple line along the margin of the petal, filamentous staminodes absent or few, white, apically purple, surrounding the white filaments in a central cone or with convex sides forming a dome, in such flowers the long tips curling during anthesis and the basally placed papillae densely interwoven, anthers yellow or purple; Fr hard, brown, long persistent on the plant, the capsule base petering out into the fruit stalk, valves with very high rims

H.E.K. Hartmann (*) Biozentrum Klein Flottbek, Hamburg, Germany e-mail: [email protected]

forming a high top, opening into an erect position, not bending backwards, expanding keels from a sharp inner edge merging into expanding sheets, extending only a little into the locules, valve wings absent, covering membranes resilient, stout, with a low yet well-developed closing ledge below the distal rim, closing body as seen from above shaped like a lens in side-view, the long axis of the lens pointing to the centre, white, splitting easily, when pressed, into two halves between which the upper end of the placenta is squeezed, narrow central column present, locules 5; S dark brown, testa with well developed, distinct and separated naps in rows; Chr 2n = 18; Ecol in Renosterbosveld and Fynbos, >200 mm rainfall p.a. in winter or at all seasons (Fig. 1); Distr SW part of WC, S Africa. Note: Known in Europe since the beginning of the eighteenth century, the genus has been treated in a revision (Glen 1986), including several species kept in Ruschia until then. The compact growth with a thick tap-root and thick, mostly short rhizomes and the distinct vittate flowers on long stalks were considered to separate the species now held in Acrodon clearly from all species of the predominantly shrubby plants of the genus Ruschia. In view of the inclusion of two procumbent species at this date, such distinction cannot be upheld. Hartmann (1996) suggested that the lens-shaped closing bodies splitting lengthwise upon pressure may be a synapomorphy of the genus, a feature that still needs to be demonstrated for the two procumbent species. Based on growth

# Springer-Verlag GmbH Germany 2017 H.E.K. Hartmann (ed.), Illustrated Handbook of Succulent Plants: Aizoaceae, DOI 10.1007/978-3-662-49260-4_2

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Fig. 1 Distribution. Members of the genus occur in the southern winter rainfall region and also in the area expecting rain at all seasons (Copyright H.E.K. Hartmann) Key to the Species of Acrodon 1. – 2. –

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Plants procumbent but not creeping, with long, trailing stems Plants compact or caespitose, developing a short, thickish rhizome with age Leaves trigonous, slender, 12–22 mm long, about 3 mm thick and wide, 0–3 teeth along keel, fewer on margins Leaves triquetrous, 5–20 mm long, about 6 mm thick and wide, 3–6 teeth along keel, fewer along the sharp margins, sometimes also some on the sides of the leaves, and sometimes none visible, but with only raised portions of the margins and keel Plants caespitose, to 60 cm in diameter and up to 30 cm tall, bracteoles embracing the calyx base Plants compact to very shortly branched, bracteoles not reaching the calyx base Leaves stout: length : width < thickness Leaves slender: length : width > thickness

and leaves, species of Acrodon have been confused with members of the genus Brianhuntleya. The distinction between the two genera lies in the fruit morphology mainly, the latter genus developing tumble fruits, a feature absent in Acrodon (Hartmann & Niesler 2012). In addition, a new species is described, based on the lens-shaped closing bodies mainly.

2 3 A. parvifolius A. deminutus

A. caespitosus 4 A. bellidiflorus A. subulatus

A. bellidiflorus (L.) N.E.Br. GC 1927: 12
M. bellidiflorum L. 1753: 482
M. bellidiflorum L. var. glaucescens Haw. 1821: 106 LT designated by Glen 1986: 214 as iconotype, Dillenius 1732: t. 189, f 233 = A. bellidiflorus (L.) N.E.Br. var. striatus (Haw.) N.E.Br. JB 1928: 78
M. bellidiflorum L. var. striatum Haw. 1795: 143, 147 T ? = A. bellidiflorus (L.) N.E.Br. var. viridis

Acrodon RUSCHIOIDEAE

(Haw.) N.E.Br. JB 1928: 78
M. bellidiflorum L. var. viride Haw. 1821: 106 NT, designated by Hartmann 2001: 24, Salm Dyck 1840: § 12, f 1β = Ruschia constricta L.Bolus NM2 1935: 496 T Bolus 718/33 (BOL!) = A. duplessiae (L. Bolus) Glen 1986: 217
Ruschia duplessiae L. Bolus NM2 1934: 431 T du Plessis 1048/32 (BOL!) non Bolus 21425, as cited with the original description = Ruschia longifolia L.Bolus NM2 1935: 500
Ruschia graminea H.Jacobsen 1955: 1640, nomen illeg. T Herre 10324 (BOL!) = Ruschia macrophylla L.Bolus NM2 1932: 351 T Compton 1188/24 (BOL!). Compact with a taproot, rarely with some short internodes in few populations; L dark green, 3–4 broad, flexible teeth on keel and margins, or margins smooth, the upper leaf surface narrowing from a broader base to a pointed tip, lower side of leaf often a little widened below the tip, L 20–60 (100) mm l, 7–9 mm b and t; Fl 1, bracteoles basal, sometimes reaching the middle of the stalk, calyx lobes a little unequal, the longest not as long as the petals which are white with a purplish-red margin and central stripe, 0–36 white, apically reddish filamentous staminodes in a short cone, anthers purple or yellow; Fr on persistent erect stalks, 30–50 mm total length, ∅ 10–12 mm; S with distant naps in rows, 1–1.25 mm l, 0.8–1 mm b, rarely bigger; Ecol on gravelly soils in open patches of shrubby associations, also often in grass and partially shaded, winter rainfall or rain at all seasons; Distr in a belt from 19 300 to 22 20 E, WC, S Africa. Note: The species shows the widest distribution area of all in the genus and occurs in most diverse habitats. It is most similar to A. subulatus but differs in having stouter leaves (Hartmann 1996). The name Ruschia longifolia has also been used for a species of Astridia, and some confusion has been caused by misapplications: this new name is a superfluous substitute for the legitimate name Ruschia longifolia L. Bolus NM2 1935. The citation of Ruschia jacobseniana in this place is an error because that name was given as a substitute for the younger homonym Ruschia longifolia (L.Bolus) L. Bolus NM3 1950
Astridia longifolia. The

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Fig. 2 Acrodon caespitosus. The very long bracteoles embrace the calyx base, a unique feature in the genus (Copyright I.N. Niesler)

mistake has been repeated by Bolus JSAB 1961: 170. A. bellidiflorus var. striatus = A. bellidiflorus A. bellidiflorus var. viridis = A. bellidiflorus Acrodon caespitosus H.E.K.Hartmann in Hartmann & Niesler 2012: 51 T Hartmann & Bayer 34493 (HBG). Plants forming cushion-shaped big and low shrubs, with 60 cm in diameter and a height of 30 cm the biggest in the genus. I spongy with a smooth surface, orange and with two lateral folds. L about 50 mm long, 5–9 mm broad and thick, the biggest in the genus, 3–12 keel teeth, only 1–2 along margins. Fl 1, bracteoles reaching up to the calyx and embracing its base, but starting from near the base of the stalk (Fig. 2), the calyx lobes overtopping the petals in the young flower, the flower itself of the short cone type, like those of A. bellidiflorus and A. subulatus, the central cone of about 300 stamens with only few or no filamentous staminodes that are equipped with long basal papillae pointing inwards forming an

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Fig. 3 Acrodon deminutus. A plant in rich fruit; note that the three capsules of one inflorescence often sit in a row (Copyright M.B. Bayer)

interwoven layer like a thick ring of felt; the 45–55 petals vittate and with marginal purple margins. Fr borne on long stalks of about 85 mm above the plant, their stalks bent upwards, the capsule proper the biggest in the genus with diameters and heights of about 12 mm, the top with 5 mm exceeding the sizes of all other species; Ecol on coastal lime stone or sandy patches in open spaces among shrubby coastal Fynbos with many invaders like Acacia cyclops or Acacia longifolia, expecting rainfall at all seasons; Distr in the lower ranges of the Breede River near its mouth, Swellendam, WC, South Africa. Note: The species has been collected earlier by different people, but a description was only offered recently by Hartmann (Hartmann & Niesler 2012: 51), based on ample living material and studies in habitat. A. deminutus Klak 2003: 118 T Bruyns 6854 (BOL) Some decumbent branches from a denser centre, up to 50 cm in diameter (Fig. 3), the internodes at first distinctly flattened, later roundish, orange, later grey, up to 15 mm long; L triquetrous with teeth along keel and margins, sometimes also on the sides, 5–10 mm long, 3.5–6 mm wide,

H.E.K. Hartmann

5–6 mm thick; Fl solitary or ternate, bracteoles embracing the short stalk completely, petals magenta or white with magenta bases and tips, in all cases vittate, the filamentous staminodes distinctly longer than the stamens, curling their ends during anthesis, magenta coloured and with long papillae pointing inwards, interwoven and forming a dense felted layer that remains visible between the thin purple filamentous staminodes themselves, the stamens also magenta; Fr usually in threes, more rarely solitary, the lens-shaped closing bodies not splitting readily upon pressure, reaching the distal recurving of the covering membranes, blocking the opening completely, diameter 6–8 mm; S with naps and about 1 mm long; Ecol in open spaces on clayey silt or among an almost paved stony surface, occurring in the west of the all-season-rainfall area; Distr known from a rather small region between 20 250 and 20 350 east and around 34 300 S, Swellendam, WC, S Africa. Note: The vittate flowers of this species differ from the most common short cone type in expressing looser filamentous staminodes with curly tips overtopping the stamens. The closing bodies are broader than the typical form and do not split readily upon pressure, similar to the situation in A. parvifolius. A. duplessiae = A. bellidiflorus A. leptophyllus = A. subulatus A. parvifolius du Plessis NM3 1958: 386 T du Plessis 164 (BOL!). Procumbent with long, trailing branches, I visible, round, grey, 6–15 mm l, ∅ up to 10 mm; L trigonous, bright green or purplish suffused, very slender,  20 mm l, 3–4 mm b and t, 0–4 low, broad-based teeth along keel, margins also smooth; Fl with basal bracteoles embracing the calyx base, petals magenta or purple, vittate and purple along margins, many purple filamentous staminodes present, forming a convex dome, with white tips curling with age and with many white long interwoven papillae towards the interior, visible to the naked eye, forming a felt, filaments rich pink, anthers pink;

Acrodon RUSCHIOIDEAE

Fig. 4 Acrodon parvifolius. An open capsule with its valves in the typically erect position; note that the closing bodies possess apical-dorsal folds, reminding of the two halves into which closing bodies of other species can split (Copyright H.E.K. Hartmann)

Fr 1, base petering out into the stalk, closing body with a distinct apical fold (Fig. 4), but not splitting easily, ∅ ca. 8 mm, ca. 20 mm l; S with distant naps in rows, ca 1.1 mm l, 0.8 mm b; Ecol on clayey river banks; Distr Hermanus, WC, S Africa. Note: One of the two species of Acrodon forming a ground cover, best recognized by its small, slender trigonous leaves, its flowers not forming a low cone type but urn-shapes like in the genus Amphibolia, the tips turning abruptly upwards and curling with age, and its rather small seeds with well-spaced naps. A. purpureostylus
Brianhuntleya purpureostyla A. quarcicola
Brianhuntleya quarcicola A. subulatus (Miller) N.E.Br. JB 1928: 77
M. subulatum Miller 1768: n 10
M. bellidiflorum L. var. subulatum (Miller) Haw. 1821: 106 NT, designated by Hartmann 2001: 25, Salm-Dyck 1840: § 12 f 1γ = M. simplex de Candolle 1800: t. 41
M. bellidiflorum L. var. simplex (DC.) Salm-Dyck 1840: § 12, text for f 1γ LT, designated by Hartmann in Hartmann & Niesler 2012: 53, de Candolle 1800: t. 41 = A. leptophyllus (L.Bolus) Glen 1986: 217
Ruschia leptophylla L.Bolus NM2 1932: 333 T de Villiers 958/31 (BOL!).

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Compact with a taproot; L subulate, i.e. tapering from a broader base to a pointed tip, the lower leaf surface narrowing acropetally, dark green, few broad teeth near the apex on keel and margins, sometimes absent in parts, L 35–70 mm l, 3–5 mm b and t; Fl with basal bracteoles, not reaching the middle of the stalk, petals white with purple margins and central line, i.e. vittate, the stamens and filamentous staminodes forming a tightly kept central short cone during the entire anthesis, ∅ ca 25 mm; Fr with long erect stalks, the valves about 1/3 of the height of the narrow locule, ∅ 9–11 mm; S with distant naps in rows, >1.3 mm l, >1 mm b; Ecol on gravelly open patches in Renosterbosveld and coastal Fynbos, shaded at times, rainfall in winter only; Distr Caledon, WC, S Africa. Note: A. subulatus represents a slenderleaved, bigger seeded version of A. bellidiflorus, sharing the few broad-based teeth with the latter species. Once the complex has been investigated in more detail, including fieldwork, A. subulatus may perhaps have to be considered a subspecies of A. bellidiflorus. The illegitimate name M. simplex has only recently been discovered again (Hartmann & Niesler 2012).

References Bolus L. (1928–1935) Notes on Mesembrianthemum and allied genera. (NM) 2: 1–508. 1–16 (9.11.1928), 17–32 (21.12.1928), 33–48 (24.1.1929), 49–64 (12.4.1929), 65–80 (3.5.1929), 81–94 (6.6.1929), 95–110 (4.7.1929), 111–129 (16.8.1929), 130–146 (4.11.1929), 147–160 (22.11.1929: the last two texts end in the middle of a description), 161–176 (20.2.1930), 177–192 (9.5.1930), 193–208 (15.8.1930), 209–224 (12.11.1930), 225–244 (12.2.1931), 245–268 (1.5.1931), 269–292 (3.7.1931), 293–308 (24.9.1931), 309–336 (29.1.1932), 337–356 (24.6.1932), 357–376 (5.12.1932), 377–396 (19.5.1933), 397–416 (16.10.1933), 417–436 (26.1.1934), 437–452 (23.5.1934), 453–472 (17.8.1934), 473–508 (11.2.1935). 517–520 Corrigenda. University of Cape Town. Bolus Herbarium. Cape Town Bolus L. (1936–1958) Notes on Mesembryanthemum and allied genera. NM3: 1–19 (5.2.1936), 21–44

30 (31.7.1936), 45–78 (14.1.1937), 79–114 (12.8.1937), 115–138 (24.3.1938), 139–188 (20.7.1939), 189–236 (25.8.1950), 237–288 (30.8.1954), 289–417 (30.4.1958). University of Cape Town. Cape Town Bolus L. (1961) Notes on Mesembryanthemum and allied genera. JSAB 27: 111–123, 169–180, 255–265 Brown N.E. (1927) Mesembryanthemum . . . GC 81: 12, 31–33, 52–53, 70, 84–85, 113–114, 116, 251–253, 430; GC 82: 92–93, 113–114, 228, 251, 263, 290–291, 348–349, 369, 390, 409, 429–430, 449–450, 470–471, 489–490, 509–511 Brown N.E. (1928) Mesembryanthemum and allied genera. JB 66: 75–80, 106–107, 138–145, 171–172, 267–268, 322–327 Candolle A.P. de (1800–1804) Plantes grasses. t. 26, 41, 53 (1800); 60 (1800 or 1801); 83, 84 (1801); 89 (1801 or 1802); 95 (1802); 129, 130 (1803); 147 (1804). Garnery, Paris (all dates after Rowley, G.D. 1956, 1957) Dillenius J.J. (1732) Hortus elthamensis . . . The author, London Glen H.F. (1986) Numerical taxonomic studies in the subtribe Ruschiinae (Mesembryanthemaceae) – Astridia, Acrodon and Ebracteola. Bothalia 16: 203–226

H.E.K. Hartmann Hartmann H.E.K. (1996) Miscellaneous taxonomic notes on Aizoaceae. Bradleya 14: 29–56 Hartmann H.E.K. (2001) Illustrated Handbook of Succulent Plants: Aizoaceae A-E. Springer. Heidelberg Hartmann H.E.K., Niesler I. (2012) A Gordion knot in Ruschioideae (Aizoaceae): Acrodon and Brianhuntleya in an attempt to disentangle the confusion. Bradleya 30: 33–60 Haworth A.H. (1795) Observations on the genus Mesembryanthemum . . ., part 2. The author, London Haworth A.H. (1821) Revisiones plantarum succulentarum. R. & A. Taylor, London Jacobsen H. (1955) Handbuch der sukkulenten Pflanzen 3. Fischer, Jena Klak C. (2003) New combinations, a new genus and five new species in the Aizoaceae. Bradleya 21: 107–120 Linnaeus C. von (1753) Species plantarum. Facs. 1957. Ray Society, London Miller P. (1768) Gardeners' dictionary ed.8: Mesembryanthemum: no. 1–32. Trianthema: no. 1–2. Acrodon: n 10. The author, London Plessis, R. du (1958) Acrodon parvifolia. NM3 1958: 386. Salm-Reifferscheidt-Dyck J. Duke of (1836–63) Monographia generum Aloe et Mesembryanthemi. I–V: Arnz, D€ usseldorf; V,VI: Henry & Cohen, Bonn. VII: M. Cohen & fil., Bonn

Acrosanthes AIZOOIDEAE H. E. K. Hartmann

Acrosanthes Ecklon & Zeyher 1837: 328; Adamson 1959: 23–28; Hartmann 2001: 25–26 Lectotypus, designated by Adamson 1959: 23 as “type species”, A. anceps (Thunberg) Sonder = Aizoon L. subg. Acrosanthes (Ecklon & Zeyher) D.Dietrich 1842: 130 LT, designated by Hartmann 2001: 25, Aizoon fistulosum (Ecklon & Zeyher) D. Dietrich  A. fistulosa Ecklon & Zeyher = A. anceps (Thunberg) Sonder Etym Gk akros, point, top, apex anthos, flower, referring to the terminal flowers above a forking = Didaste E. Meyer ex Sonder in Harvey & Sonder 1862: 472 nomen nudum. Mat-forming or sprawling with long branches, I woody, at least at their bases; L terete to flat, glabrous; Fl solitary, seemingly lateral, but in reality terminal and overtopped by one of the side branches developing below, perigon 5-lobed, lobes shorter than the basal tube, white inside, stamens 8-many, mostly in groups, ovary perigynous, divided basally only incompletely by low septa into two locules, a single ovule per locule in basal position; Fr a xerochastic capsule with a parchment-like wall persisting in the drying perigon; S compressed and reniform or orbicular, testa rugose; Ecol winter rainfall >400 mm p.a. (Fig. 1); Distr W districts of WC, S Africa.

H.E.K. Hartmann (*) Biozentrum Klein Flottbek, Hamburg, Germany e-mail: [email protected]

Note: The genus has received little attention since its establishment in 1837, hence no photographs are available; instead, drawings are presented here. The present treatment follows Adamson 1959: 23–28 essentially, including the lectotypification of Acrosanthes, but with an additional lectotypification of Aizoon subg. Acrosanthes. A new study based on fresh, growing material is highly desirable, focussing in particular on the constancy or variation in growth forms and establishing whether indeed the longitudinal slits seen in capsules on herbarium material represent a mode of opening entirely different from the circumscissile fruits in most genera of the subfamily Sesuvioideae. The epidermis of the leaves should also be examined in order to assess whether the genus does indeed belong in the Aizoaceae. On herbarium sheets, the needleshaped leaves of some species appear in a shiny olive green quite unlike any other leaves in the family. The complete lack of any expanding tissue and the basal 1–2 ovules separate the genus well from Aizoon, where it had been placed forming a subgenus (Dietrich 1842: 130, not 1847, as given by Adamson 1959: 23). The name Didaste E. Meyer appears in Sonder (Harvey & Sonder 1862: 472, 473) without any description and only in three synonyms for three species of Acrosanthes, excluding the type species of the latter genus, A. anceps. A. anceps (Thunberg) Sonder in Harvey & Sonder 1862: 472  Trianthema anceps Thunberg 1794:

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Leaves terete, stamens 25 and more Leaves flat, stamens always 25, not in groups, filaments dark, anthers white (Fig. 5); Fr rounded, shorter than the perigon; S dark brown, rugose; Ecol sandy or stony flats and lower slopes; Distr from Piquetberg to Bredasdorp, WC, S Africa. Note: The species differs from all others of the genus in the high number of stamens never arranged in groups and in the terete leaves.

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lacks any description and is, therefore, a nomen nudum. D. decandra = Acrosanthes angustifolia D. pentandra = Acrosanthes humifusa D. icosandra = Acrosanthes teretifolia

References

Fig. 5 Acrosanthes teretifolia, drawn from a sheet of the isotype of Ecklon & Zeyher 2148 kept at OXF, showing a virtual longitudinal section through the flower with the dome-shaped gynoecium half as high as the tepals and two of the five distinct bundles of stamens between them, each with five elements; drawn with permission (Copyright H.E.K. Hartmann)

Didaste = Acrosanthes The name Didaste E.Meyer was used by Sonder (in Harvey & Sonder 1862: 472–473) first, who listed three species as synonyms in the genus Acrosanthes, attributing the names to E. Meyer, who had probably named sheets collected by Drege. Didaste E.Meyer ex Sonder

Adamson R.S. (1959) The South African species of Aizoaceae. JSAB 25: VI. Acrosanthes: 23–28 Dietrich D. (1842) (neither 1843 nor 1847, as cited elsewhere): Synopsis plantarum . . .: Aizoon: 128–130. Voigt. Weimar Ecklon C.F., Zeyher C. (1837) Enumeratio plantarum africae australis extratropicae . . .3: Ficoideae: 308–329. The authors, Hamburg Fenzl E. (1839) Monographie der Mollugineen. Annalen des Wiener Museums der Naturgeschichte 2: 243–310 Hartmann H.E.K. (2001): Illustrated Handbook of Succulent Plants: Aizoaceae A-E. Springer. Heidelberg Harvey W.H., Sonder W. (1862) Flora Capensis 2: 386–478, 596–598, 616. Hodges, Smith & Co, Dublin Sonder O.W. (1862) Aizoon. Galenia. Mesembryaceae. Trianthema. In: Harvey W.H., Sonder W. (eds) Flora Capensis Thunberg P. (1794) Prodromus plantarum capensium, quas in Promontorio Bonae Spei Africae annis 1772–1775 collegit Pars prior: 80. Edman, Uppsala

Aethephyllum RUSCHIOIDEAE H. E. K. Hartmann

Aethephyllum N.E.Br. MDG 1928: 400; Ihlenfeldt & Struck 1987: 411–453; Hartmann 2001: 26–27; Klak & Bruyns 2012: 293–307; Klak 2012: 27–33  Micropterum Schwantes subg. Aethephyllum (N.E.Br.) Schwantes KuaS 1950: 7 Typus A. pinnatifidum (L.f.) N.E.Br.  Micropterum pinnatifidum (L.f.) Schwantes Etym Gk aethe, irregular, unusual, phyllon, leaf, referring to the lyrate or pinnatifid leaves. Short-lived annuals with decumbent branches; L pinnatifid, bifacial, epidermis with prominent bladder cells, wax cover thin, rugose; Fl apparently solitary, but in reality almost the whole plant representing the dichasial Inf, pedicellate, petals white or yellow, stamens yellow with darker anthers, filaments basally papillate, 5 flat nectaries; Fr pedicels s-shaped, base of capsule short funnel-shaped, top flat, covering membranes short, translucent and flexible, expanding sheets rising into high rims and merging distally (in the open capsule) into a thin, flimsy layer, no distinct expanding keel discernible, valve wings broad over the entire length of the valve, 5 locules; S testa with distant naps; Ecol plentiful after fires, mostly in Fynbos, but also coastal, winter rainfall (Fig. 1); Distr from Kamieskroon, NC,

H.E.K. Hartmann (*) Biozentrum Klein Flottbek, Hamburg, Germany e-mail: [email protected]

over Clanwilliam, Piquetberg, Tulbagh, Vanrhynsdorp, to the Cape, WC, S Africa. Note: Based on more recent studies in the Dorotheantheae (Klak & Bruyns 2012), the genus Aethephyllum is enlarged here in order to include three species with lyrate leaves (see also key in Klak 2012: 32), requiring two new combinations. Aethephyllum herrei (Schwantes) H.E.K. Hartmann comb. nov.  Micropterum herrei Schwantes Kakteen und andere Sukkulenten 1, 1950: 8 as basionym = Cleretum herrei (Schwantes) Ihlenf. & Struck 1987: 437 LT, designated by Ihlenfeldt & Struck 1987: 437, Herre s.n. 1949 (HBG). Ascending in growth, rather robust, I 3.5 mm ∅; L lyrate, 40 mm l, 15 mm b, epidermis with big bladder cells; Fl  8 mm ∅, sessile, 6–9 white to purple petals, 6–8 filamentous staminodes, 15–17 stamens basally papillate, filaments yellowish, anthers orange (Fig. 2); Fr stalks s-shaped, lateral rims instead of covering membranes; S roundish in side view, testa cells convex with undulate anticlinal walls, greyish-black; Ecol sandy places, near the sea shore, fire successor; Distr Cape Town to Simonstown, Somerset West, WC, S Africa Note: Morphologically, A. herrei differs from A. pinnatifidum in its more robust habit, reduced covering membranes, and convex testa cells without naps.

# Springer-Verlag GmbH Germany 2017 H.E.K. Hartmann (ed.), Illustrated Handbook of Succulent Plants: Aizoaceae, DOI 10.1007/978-3-662-49260-4_4

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B.E.

Mi

Um

Cr

G.R.

St

Gh

E.L.

Ui P.E.

All seasons

Fig. 1 Distribution of Aethephyllum is restricted to the southern part of the winter rainfall area (Copyright H.E.K. Hartmann) Key to the Species of Aethephyllum 1. – 2. –

Petals yellow, top of capsules flat Petals white to purplish suffused, capsule tops with ribs Fruit stalks short and s-shaped, seeds greyish-black (anthracite) Fruit stalks 30–40 mm long, almost straight, seeds brown

Fig. 2 Aethephyllum herrei in flower after rains in August 2012; note the few and short petals and the prominent erect valve rims in the ripening central capsule (Copyright C. Wakfer)

A. pinnatifidum 2 A. herrei A. lyratifolium

Aethephyllum lyratifolium (Ihlenf. & Struck) H. E.K.Hartmann comb. nov.  Cleretum lyratifolium Ihlenf. & Struck Beiträge zur Biologie der Pflanzen 61, 1987: 437 as basionym, Typus Bayer 2596 (HBG). Decumbent with gross bladder cells; L irregularly lyrate, amplexicaul; Fl stalks short or absent, 4–5 mm ∅, petals ca. 1 mm l, white, stamens 5; Fr stalks 30–40 mm l, base roundish, low, top with low valve rims, covering membranes absent to half over the locules, ∅ 8–10 mm; S triangular in side view, brown, testa cells concave, anticlinal walls straight; Ecol in gravel; Distr from Khamieskroon, NC, to Laingsburg, WC, S Africa. Note: This species is most similar to A. herrei, from which it differs in the straight fruit stalks, the

Aethephyllum RUSCHIOIDEAE

low valve rims, and the brown seeds, as opposed to blackish-grey ones in A. herrei. A. pinnatifidum (L.f.) N.E.Br. MDG 1928: 400  M. pinnatifidum L.f. 1781: 260  Cleretum pinnatifidum (L.f.) L.Bolus SAG 1927: 327  Cleretum pinnatifidum (L.f.) N.E.Br. GC 1927: 228 nomen illeg.  Micropterum pinnatifidum (L.f.) Schwantes KuaS 1950: 7 LT, designated by Hartmann 2001: 27, “M. pinnatum” 649.7 (LINN) = M. pinnatum Thunberg 1791: 15 LT, designated by Hartmann 2001: 27, Thunberg 12003 (UPS). L at first opposite, later on all flowering branches spirally arranged; Fl pedicels elongating before the buds open, papillae on filaments as thin threads; S almost round in side view, the naps well separated; Ecol in coastal Fynbos, fire successors; Distr from Vanrhynsdorp to Tulbagh, WC, S Africa. Note: A. pinnatifidum differs from the other two species in its yellow petals and the absence of ribs on the valves. As in other annuals, seeds from habitat germinate tardily in cultivation, requiring treatment with KOH or some mechanical destruction of the seed coat to improve germination success.

39

References Bolus L. (1927) South African succulents . . .: SAG 17: 134, 238–239, 256, 280–281, 326–327, 365, 398–400, 433–435 Brown N.E. (1927) Mesembryanthemum . . . GC 81: 12, 31–33, 52–53, 70, 84–85, 113–114, 116, 251–253, 430; GC 82: 92–93, 113–114, 228, 251, 263, 290–291, 348–349, 369, 390, 409, 429–430, 449–450, 470–471, 489–490, 509–511 Brown N.E. (1928) Ergänzungen zu dem MesembriaceenAufsatz. MDG 43: 400–401 Hartmann H.E.K. (2001): Illustrated Handbook of Succulent Plants: Aizoaceae A-E. Springer. Heidelberg Ihlenfeldt H.-D., Struck M. (1987) Morphologie und Taxonomie der Dorotheanthinae Schwantes (Mesembryanthemaceae). BBP 61: 411–453 Klak C. (2012) Two new species of Cleretum (Ruschioideae-Dorotheantheae, Aizoaceae) from the Northern Cape . . . . Haseltonia 18: 29–35 Klak C., Bruyns P.V. (2012) Phylogeny of the Dorotheantheae (Aizoaceae). . . . Taxon: 61, 293–307 Linnaeus C. fil. (1781) Supplementum plantarum. Systematis vegetabilium – ed. 13 generum plantarum: 227, 258–261, 392. At the expenses of the orphanage, Braunschweig Schwantes G. (1950) Micropterum Schwantes. KuaS 1(2): 4–5, (3) 6–8 Thunberg P. (1791) Descriptiones Mesembryanthemorum quorundam in Capitis Bonae Spei Africes . . . Novorum Actorum medico-physicorum academiae Caesareae Leopoldino-Carolinae naturae curiosorum in Germania 8: App.: 1–18

Aizoanthemum AIZOOIDEAE H. E. K. Hartmann

Aizoanthemum Dinter ex Friedrich 1957: 342  Aizoanthemum Dinter Kakteenkunde 1935: 27 nomen invalid.  Aizoon L. subg. Aizoanthemum (Friedrich) Adamson 1959: 44; Hartmann 2001: 28–29 Typus A. membrumconnectens Dinter ex Friedrich  Aizoon membrumconnectens (Dinter ex Friedrich) Adamson = A. rehmannii (Schinz) H.E.K.Hartmann Etym Aizoon, a genus of Aizoaceae, Gk anthos, flower, the name stressing the similarity to the genus Aizoon. Annual herbs, with erect to spreading, rarely decumbent branches, I young with longish papillae, old ones with distant, shorter papillae, appearing as flat scales when dry; L alternate, only in the distal part of the Inf opposite in some cases, elongateoval, often narrowing towards the base and sometimes developing a petiole, papillae low and inconspicuous or more prominent; Fl in large Inf comprising  the entire plant, mostly purely dichasial in construction, perigynous, perigon papillate outside, greenish, white, or yellow inside, all lobes of  equal length, stamens numerous, basally flat; Fr  globose or columnar, the valves opening completely by parallel, long expanding keels reaching from the centre of the fruit to the tip of the valve, 5–10 locules; S reniform to round, with concentric ridges, dark brown to black; Ecol in

H.E.K. Hartmann (*) Biozentrum Klein Flottbek, Hamburg, Germany e-mail: [email protected]

sandy soils, often near the coast (Fig. 1); Distr S Angola, N Namibia, from the Canary Islands and S Spain along the S coasts of the Mediterranean Sea to Syria, Armenia, Iraq, and Iran. Note: The genus name was first presented by Dinter (1935: 27) without a Latin description, it was therefore invalidly published (ICN McNeill et al. 2012: Art. 39). Friedrich (1957: 342) established the genus based on an extensive discussion of the discriminating features; his view is followed here; the placement of the taxon as a subgenus in Aizoon (Adamson 1959: 44) is not upheld. As diagnostic character states are seen the basally flat filaments and the hygroscopic capsules, opening completely by extensive expanding keels, both features absent in Aizoon. In this sense, the genus comprises 5 species. A. bossii = A. dinteri A. dinteri (Schinz) Friedrich 1957: 344  Aizoon dinteri Schinz 1898: 523 T Dinter 55 (Z) = A. bossii Dinter ex Friedrich 1957: 344 nomen nudum = A. sphingis Dinter ex Friedrich 1957: 344 nomen nudum = A. stellatum Dinter ex Friedrich 1957: 344 nomen nudum. Erect, highly branched, 20–40 cm h; L sessile to petiolate near the base of the plant, ovate to obovate, obtuse, finely papillate, 10–60 mm l, 7–35 mm b; Fl  sessile, perigon 5–6 mm l, lobes spreading with age, greenish inside, c. 50 stamens, filaments sharply denticulate at their

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Fig. 1 Distribution of Aizoanthemum falls into two parts: southern Africa and along the southern border of the Mediterranean Sea, extending into the Near East (Copyright H.E.K. Hartmann)

Key to the Species of Aizoanthemum 1. – 2. – 3. – 4. –

Locules 5 Locules 7–10 Leaves with a petiole c. as long as the blade Leaves narrowing  abruptly at the base, sessile and not petiolate Perigon 3–4 mm l, fruit 1.5 mm ∅ Perigon 7–15 mm l, fruit 5–6 mm ∅ Perigon 5–7 mm l, fruits of a population mostly with 7–8 locules Perigon 8–10 mm l, fruits of a population mostly with 10 locules

edges, the outer ones sometimes sterile; Fr subglobose, 3–3.5 mm ∅, mostly 7–8 locules; Ecol in sandy or brackish soils; Distr from Kunene (Kaokoveld) to Grootfontein Dist., Otjozondjupa, and Swakopmund Dist., Erongo, Namibia. Note: The species is similar to A. rehmannii in the high number of locules per fruit, it differs in the more erect growth and the smaller flowers and fruits. A. galenioides (Fenzl ex Sonder) Friedrich 1957: 344  Aizoon galenioides Fenzl ex Sonder in Harvey & Sonder 1862: 469 T Drege 7060 (W, lost), NT, designated by Friedrich 1957: 344, Dinter 7884 (M).

2 4 A. mossamedense 3 A. galenioides A. hispanicum A. dinteri A. rehmannii

Erect, highly branched, to 15 cm h; L with a short petiole, oblanceolate to obovate, papillate, 7–20 mm l, 3–7 mm b; Fl sessile, perigon 3–4 mm l, lobes erect, cucullate, stamens 25–30, filaments with smooth edges; Fr globose, 1.5 mm ∅, 5 locules; S c. 0.5 mm b; Ecol in coastal sands; Distr Omaruru Dist., Swakopmund Dist., Erongo, Namibia. Note: The species can be recognized best by the tiny, globose, 5-locular fruits. A. hispanicum (L.) H.E.K. Hartmann 2001: 29  Aizoon hispanicum L. Species plantarum 1753: 488  Aizoon hispanicum L. var. genuinum Maire 1962: 279 nomen illeg. LT, designated by

Aizoanthemum AIZOOIDEAE

Fig. 2 Aizoanthemum hispanicum in flower in habitat in Ein Gedi, Israel, in December 2011; note the irregular numbers of stamens in the groups (Copyright C. Barnick)

Peruzzi et al. 2004: 541, Dillenius 1732: t.117 f 143 = Aizoon hispanicum L. var. minus Maire 1962: 280 nomen invalid. T Joly s.n. (?) = Aizoon sessiliflorum Moench 1794: 698 T? Erect, to 25 cm h, consisting of a highly branched terminal cyme and 2–3 basal additional ones, I with  dense, nearly globose papillae; L sessile, basally opposite, alternate in the middle and opposite again in upper parts of the cymes, covered by shining, low, roundish papillae, 5–40 mm l, 1–9 mm b; Fl in rich cymes,  sessile, perigon with vesicular papillae outside, white inside, the lobes prominently keeled from protruding bundles, 4–9 mm l, stamens in five groups (Fig. 2); Fr a five-angled column with 5 prominent distal corners giving it a star-shaped appearance when seen from above, opening completely by long expanding keels, ∅ 5–6 mm, 5 locules; S black, 0.75–1 mm l; Chr 2n = 32; Ecol in sandy to slightly saline soils in predominantly Mediterranean climates, also in fine gravel; Distr Algeria; Armenia; Egypt; Iran; Iraq; Israel; Italy; Libya; Morocco; Portugal (Madeira: Porto Santo); Spain, including Gran Canaria and the southern mainland; Syria; Tunisia. Note: The data collected from floras show a wide distribution range for the species, similar to that of Aizoon canariense, but extending further into Asia. Of the two varieties distinguished by Maire (1962: 279–280), one is illegitimate because it is based on the type of the species, the other is invalid because

43

Fig. 3 Aizoanthemum mossamedense in flower with many stamens and long papillae on the outside of the perigon (Copyright N. J€ urgens)

it lacks a Latin diagnosis or description. Aizoon sessiliflorum was explicitly named for Aizoon hispanicum, but the type is not known. The lectotypification of Peruzzi et al. (2004) is based on the fact that the sheet LINN 650.2 (LINN) considered to be the lectotype by Hartmann 2001: 29 does not bear an annotation on it indicating that Linnaeus saw it before publishing that name. Hence, another lectotype had to be chosen, and the only reference to a figure that is available is found in Dillenius (1732). The application of the name Aizoon hispanicum refers to the original description only and is not applicable, as discussed, e.g., in Hartmann 2001: 28–29. A. membrumconnectens = A. rehmannii A. mossamedense (Welwitsch in Oliver) Friedrich 1960: 567  Aizoon mossamedense Welwitsch in Oliver 1871: 584 T Welwitsch 1264 (BM). Diffusely branched,  erect, with some alternate leaves below the terminal, rich cyme constituting most of the plant, branches to 25 cm l, papillate to hairy; L petioles c. as long as the blade, oval to broadly elliptical, obtuse, with lower papillae than I, 17–31 mm l, 3–13 mm b; Fl sessile, perigon with long papillae outside, bright yellow inside, lobes ovate to lanceolate, stamens numerous, in a ring, filaments laterally finely denticulate (Fig. 3); Fr globose, 5 locules; S ridged; Ecol in sand near the coast; Distr Mossamedes, Angola.

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Note: The species appears to occur in a restricted area only. It can best be distinguished from the other 5-locular species by the rather big, round fruit. A. rehmannii (Schinz) H.E.K.Hartmann 2001: 29  M. rehmannii Schinz Bulletin de l’Herbier Boissier 5, Appendix 3, 1897: 80 T Kolbe s.n. (Z, lost?) LT, designated by Hartmann 2001: 29, Rehmann s.n. (Z!) = A. membrumconnectens Dinter ex Friedrich 1957: 345  A. membrumconnectens Dinter Kakteenkunde 1935: 27 nomen invalid. T Strey 2496 (M). Prostrate, the entire plant formed by a rich cyme, I with densely placed, elongate, pointed papillae; L obovate to elliptical, often conduplicate, sessile or with a short petiole, papillate, 10–30 mm l, 5–13 mm b; Fl sessile, basal tube 6–7 mm l, numerous stamens in a ring, filaments laterally finely dentate; Fr subglobose, 10 locules, ∅ 5–6 mm; S c. 0.7 mm l; Ecol in gravelly flats; Distr Rehoboth Dist., Hardap, Swakopmund Dist., Erongo, Namibia. Note: The low, prostrate growth characterizes the species best within the genus. In the description, “Miss Kolbe” is given as the collector of the material used as type. The sheet at Z, marked as “Typus” bears a heading on the label:“Dr. A. Rehmann - Exsiccata Africae austr. 1875–1880”. It can be speculated that Schinz, when describing the species and naming it after Rehmann, knew that this collection had been given to Rehmann by Kolbe. Since there is no evidence for this speculation, the sheet Rehmann s.n. marked “Typus” (Z) has been chosen as the lectotype, based on three arguments: 1. The material can be clearly identified as being the species described by Schinz under this name, 2. The sheet has been available to Schinz when he described the species first; 3. The epithet refers to Rehmann, from whose collections the “Typus” comes. Friedrich (1970: 13) listed Aizoon rehmannii as a possible synonym under A. membrumconnectens already, yet did not make the necessary transfer, which was undertaken by Hartmann 2001: 29. A. sphingis = A. dinteri A. stellatum = A. dinteri

H.E.K. Hartmann

References Adamson R.S. (1959) The South African species of Aizoaceae VII. Aizoon. JSAB 25: 29–51 Dillenius J.J. (1732) Hortus elthamensis . . . The author, London Dinter K. (1935) Aizoanthemum membrumconnectens Dtr. n. gen. Kakteenkunde 1935: 27–28 Friedrich H.-C. (1957) Aizoanthemum Dinter ex Friedr., eine wenig beachtete Gattung der Ficoideae aus S€ udwestafrika. MBSM 2: 339–347 Friedrich H.-C. (1960) Mesembrianthemenstudien I. Beitrag zur Kenntnis der Gattungen Stoeberia Dtr. et Schw. und Ruschianthemum Friedr. gen. nov. MBSM 3: 554–567 Friedrich H.-C. (1970) Aizoaceae. In: Merxm€ uller H. (ed.) Prodromus einer Flora von S€ udwestafrika 27: 1–135 Hartmann H.E.K. (2001): Illustrated Handbook of Succulent Plants: Aizoaceae A-E. Springer. Heidelberg Harvey W.H., Sonder W. (1862) Flora Capensis 2: 386–478, 596–598, 616. Hodges, Smith & Co, Dublin Linnaeus C. (1753) Species plantarum. Facs. 1957. Ray Society, London Maire R. (1962) Flore de l’Afrique du Nord. Aizoaceae: 246–289 Merxm€ uller H. (ed.) Prodromus einer Flora von S€ udwestafrika 27: 1–135 McNeill J., Barrie F.R., Buck W.R., Demoulin V., Greuter W., Hawksworth D.L. Herendeen P.S., Knapp S., Marhold K., Prado J., Prud’homme van Reine W.F., Smith, G.F., Wiersema J.H. et al. (2012) ICN International Code of Nomenclature for algae, fungi and plants (Melbourne Code). Regnum Vegetabile 154. A.R.G. Gantner Verlag. Moench C. (1794) Methodus Plantarum horti botanici et agri Marburgensis: 698. Cotta, Marburg Oliver D. (1871) Flora of tropical Africa 2: 582–588. Reeve & Co., London Peruzzi L., Passalacqua N.G., Jarvis C.E. (2004) Lectotypification of Aizoon hispanicum, Plantago albicans, and Staphylea pinnata, names of three Linnaean species occurring in Calabria (S.Italy). Taxon 53: 540–542 Schinz H. (1897) Die Pflanzenwelt Deutsch-S€ udwestAfrikas (mit Einschluß der westlichen Calahari). Bulletin de l’Herbier Boissier 5, Appendix 3: 5–101 Schinz H. (1898) Beiträge zur Kenntnis der afrikanischen Flora (Neue Folge) VII. Mitteilungen aus dem botanischen Museum der Universität Z€ urich V. Bulletin de l’Herbier Bossier, Genève 6: 523 Sonder O.W. (1862) Aizoon. Galenia. Mesembryaceae. Trianthema. In: Harvey W.H., Sonder W. (eds) Flora Capensis Welwitsch F.M.J. (1871) In: Oliver D. (1871) Flora of tropical Africa 2

Aizoon AIZOOIDEAE H. E. K. Hartmann

Aizoon L. 1753: 488 non Hill 1756: 53; Hartmann 2001a: 30–33; 2001b: 349 Lectotypus A. canariense L.  Veslingia Heister ex Fabricius 1759: 201 no species named Etym aizoon, an evergreen plant. Annual herbs to perennial shrubs, procumbent to erect branches and woody roots, I hairy; L opposite or alternate, linear and terete to spatulate or oval, with prominent, white, long hairs beside much smaller bladder-cells (papillae); Fl in  rich cymes, perigon hairy outside, yellow, rarely pink, purple, white, or cream inside, stamens numerous, in groups or in a ring, filaments filiform; Fr a loculicidal capsule opening only a little near the centre of the fruit as if lifting the 5-partite lid, the valves being fixed to the depressed centre of the fruit by the short expanding keels, 5 locules; S with concentric ridges; Ecol in arid areas, often in sand or fine gravel (Fig. 1); Distr Northern Africa: from the Canary Islands to Sokotra and along the African Mediterranean coast, also in Spain and S Italy, N Kenya, extending to India in the east and to Afghanistan in the north in Asia; in Southern Africa, south of a line from Angola to Zimbabwe. Note: A number of species now placed in other genera of the Aizoaceae has originally been

H.E.K. Hartmann (*) Biozentrum Klein Flottbek, Hamburg, Germany e-mail: [email protected]

described in Aizoon, e.g. the Australian ones now kept in Gunniopsis (Chinnock 1983). A worldwide revision is lacking, but numerous treatments of some of the species in many floras permit the presentation of an overview. The taxonomy follows mainly Adamson 1959: 29–51 and Friedrich 1970: 13–16. A. angustifolium  Acrosanthes angustifolia A. argenteum = A. rigidum A. asbestinum Schlechter JB 1897: 280 T Marloth 2075 (B) = A. burchellii N.E.Br. KB 1908: 290 T Burchell 1734 (K!) not 1753 (K), as published by Brown. Low shrubs to 15 cm h, with very thick, persistent short stems merging into an also thick, woody rootstock, dividing near ground level into several erect, rarely more decumbent, woody, hard branches (Fig. 2); L only in the apical parts of the branches, opposite, almost pungent, covered densely by white hairs, some to many of them spreading, linear to lanceolate, 8–15 as long as wide; Fl terminal and overtopped by one of the side branches, thus the Inf monochasial, perigon 6–8 mm l, stamens in a ring; Fr 3–4 mm ∅, 5 locules, centrally depressed; S light brown, densely tuberculate; Ecol on slopes with gravel in open patches of shrubby vegetation; Distr from Outjo Dist., Kunene, to Warmbad, Karasburg Dist., Karas, Namibia; NC, S Africa.

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Fig. 1 Distribution. Aizoon occurs in the winter rainfall regions of the Old World, sparing a huge gap over the tropics and extending further in the north eastern areas as far as Afghanistan (Copyright H.E.K. Hartmann)

Note: The placement of A. burchellii as a synonym here follows Friedrich (1960: 615) who stresses the high degree of variation in several characters: the long papillae forming a pungent surface in typical forms from the Asbestos mountains merge into populations with soft hairs appressed to the leaf surfaces. Independently, narrow to wider leaf shapes can be found, and growth forms reach from erect to low-oblique branches. With his description, Brown (1908: 290) gave the collecting number Burchell 1753 and a detailed information on the locality, probably copied from Burchell. The same wording regarding the locality is written on the sheet Burchell 1734 (K!) whereas number 1753 has not been found. The collecting number has, therefore, been adjusted accordingly. A. burchellii = A. asbestinum

Fig. 2 Aizoon asbestinum in side view to show the splitting stems and roots (Copyright H.E.K. Hartmann)

A. camforosma Reichenbach ex Sprengel 1827: 195 T ? Shrubs with erect branches; L alternate, linearsubulate, glabrous; Fl pedunculate, axillary; Ecol unknown; Distr unknown, S Africa. Note: The species appears to have been overlooked in all treatments available for the

Aizoon AIZOOIDEAE

Fig. 3 Aizoon canariense: a dead plant with old open fruits after watering, exposing the short white expanding keels below the lifted valves (Copyright H.E.K. Hartmann)

genus Aizoon. The pedunculate flowers suggest that it does not belong here, but no suggestions as to its position appear to have been made. A. canariense L. 1753: 488 T LINN 650.1 (LINN) = Veslingia caulifloris Moench 1802: 299 LT, designated by Hartmann 2001a: 31, Nissolle 1730: 319, t. 13 p. 324 (not t. 19, as cited by Moench)  A. procumbens Crantz 1766: 135 T Nissolle 1730: t. 13, f 1 (as Ficoidea) = Glinus crystallinum Forskål 1775: 95  Glinus procumbens Forskål 1775: 25 ex Adamson 1959: 311 nomen nudum LT, designated by Hartmann 2001a: 31, Forskål 1775: t. 14 = M. dubium Haw. 1795: 471 non Haw. 1803: 39 nomen illeg. LT, designated by Hartmann 2001a: 31, Petiver 1703/4: t. 39 f 3 = Veslingia heisteri Fabricius ex Willdenow 1799: 1053 T ? = A. spathulatum Ecklon & Zeyher 1837: 325  A. canariense L. var. denudatum

Adamson (1959: 31) cited “Glinus procumbens Forsk. Descr. 25. t.14. 175” as a synonym for A. canariense. The name Glinus procumbens does not occur in Forskål 1775, where the genus Glinus is treated on pages 95–96, not 25, and t. 14 refers to Glinus chrystallinum (Christensen 1922: 20); the figure is chosen as the lectotype here. Glinus procumbens is not listed in Index Kewensis, hence the name used by Adamson is a nomen nudum probably established in error. 1

47

Sonder in Harvey & Sonder 1862: 470 T Ecklon & Zeyher 2128 (S). Annuals with mostly decumbent branches to 40 cm l, remaining in situ as dry skeletons for several years fixed by a woody perpendicular root; L alternate, spatulate, or oval, narrowing into a petiole 3–18 mm l, blades 8–70 mm l, 4–45 mm b; Fl in dichasial or monochasial arrangement, always secund, perigon yellow inside, stamens 12–15, in bundles; Fr persisting on the dead plants for several years, truncate, the valves remaining in function for up to 2 years (Fig. 3), then disintegrating and leaving a central hollow with a small persisting column in the cente, 5–9 mm ∅; S dark brown, with concentric ridges; Ecol often on disturbed land, but always in dry situations, in sand or gravel of washes; Distr over the entire area given for the genus. Note: This is probably the species of Aizooideae with the widest natural distribution area; it occurs mostly in regions with summer rainfall, but can also stand a certain amount of winter rain. The synonymy following Adamson (1959: 31) and Gonçalves (1978: 509) should be re-examined. A. canariense var. denudatum = A. canariense A. contaminatum = Galenia secunda A. crystallinum  Galenia crystallina A. dinteri  Aizoanthemum dinteri A. elongatum = Galenia secunda A. fistulosum = Acrosanthes anceps A. fruticosum  A. schellenbergii;  Galenia fruticosa;  Galenia affinis A. galenioides  Aizoanthemum galenioides A. giessii Friedrich 1960: 613 T Volk 12730 (M). Woody plants with a thick rootstock ending in several thick, woody decumbent branches with decumbent short shoots to 5 cm h; L in terminal groups on all shoots, elliptical, acuminate, basally tapering into a  broad petiole, covered densely by white, appressed hairs, 15–75 mm l, 5–17 mm b, the relation between length and width 4–5:1 (Fig. 4); Fl secund, perigon connate for 1/3 of its length, 10–14 mm l, with long hairs outside, glabrous and bright yellow inside, stamens numerous; Fr truncate, 3–4 mm ∅, 5 locules; Ecol on limestone; Distr

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Fig. 4 Aizoon giessii with appressed white hairs, soft to the touch, on the slender leaves (Copyright H.E.K. Hartmann)

Gibeon, Mariental Dist., Maltahöhe Dist., Rehoboth, Dist., Hardap, Windhoek Dist., Khomas, Namibia. Note: Following Friedrich (1960: 614), the species differs from the similar A. asbestinum by the strictly decumbent branches, the white, soft hairs, the broader leaves and bigger flowers. It seems also that this species prefers limestone. In view of the variation in all reported distinguishing character states, a more thorough investigation is needed in order to determine whether the rank of species is justified for this taxon. A. glabrum  Gunniopsis glabra A. glinoides L.f. 1781: 261 T Thunberg 12050 (UPS!) = A. hirsutum Ecklon & Zeyher 1837: 325 T Ecklon & Zeyher 2130 (?). Prostrate herbs, I densely covered by long, erect hairs, branches 15–60 cm l; L petiolate, broadly elliptical, acuminate or cuspidate at tips, with long, dense hairs, 15–50 mm l, 5–15 mm b; Fl perigon with prominent, erect, long hairs outside, bright yellow inside, perigon lobes 3–8 mm l, spreading widely at anthesis; Fr truncate above, the central yellow area where the stigmas have broken off conspicuously smooth and yellow, young fruits often bright maroon (Fig. 5); S black, the concentric ridges broken or replaced by dots; Ecol in dry open places; Distr EC, S Africa. Note: The cover of very long and dense hairs and the erect hairs on the outer perigon characterize this species well.

H.E.K. Hartmann

Fig. 5 Aizoon glinoides with a regular row of young capsules in bright copper colour and an old capsule of the previous year in blackish brown (Copyright H.E.K. Hartmann)

A. herniarium  Galenia herniariaefolia A. hirsutum = A. glinoides A. hispanicum  Aizoanthemum hispanicum A. hispanicum var. genuinum  Aizoanthemum hispanicum A. hispanicum var. minus = Aizoanthemum hispanicum A. karooicum Compton 1931: 283 T Compton 2966 (BOL). Woody shrubs with erect to procumbent branches 5–20 cm l, I covered with long, white, appressed hairs when young, later smooth and finally black; L alternate, crowded at the base but scattered above, elliptical to lanceolate, often folded or rolled up, 15–30 mm l, 3–5 mm b; Fl sessile, perigon 4–5 mm l, covered outside with reversed hairs, stamens c. 20 in groups; Fr flattened; Ecol in sandy depressions; Distr Calitzdorp, Heidelberg, Laingsburg, Robertson, WC, S Africa. Note: The black basal internodes appear distinctive of this species. A. kochii  Gunniopsis kochii A. microphyllum = Galenia herniariaefolia A. montevidense = Sesuvium parviflorum A. mossamedense  Aizoanthemum mossamedense

Aizoon AIZOOIDEAE

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A. propinquum = Galenia secunda A. pubescens  Galenia pubescens A. quadrifidum  Gunniopsis quadrifida A. rarum = A. paniculatum

Fig. 6 Aizoon rigidum with a more irregular arrangement of the hairy flowers and with shorter and narrower leaves (Copyright H.E.K. Hartmann)

A. paniculatum L. 1753: 488 T LINN 650.6 (LINN) = A. lanceolatum L. in Murray 1774: 392 T ? = A. rarum N.E.Br. KB 1908: 291 T Burchell 1013 (K!) = A. stellatum Lamarck 1792: 418 T Sonnerat s.n. (P?) = A. tomentosum Lamarck 1792: 418 T Sonnerat s.n. (P?). Prostrate, sprawling to semi-erect shrubs with woody bases and annual, flowering branches which die back, I covered with white, loosely appressed hairs, some with two spreading arms; L opposite, alternate in the Inf, crowded at base, smaller and dispersed above, petiolate, elliptical, obtuse, some long hairs dispersed over them, 15–60 mm l, 2–6 mm b; Fl mostly in terminal monochasia appearing as a panicle, sessile, perigon 4–8 mm l, with spreading, armed hairs outside, red or pink, less often yellowish or creamcoloured inside, stamens in groups of 3–10; Fr 2–3 mm ∅; Ecol in sand; Distr western districts of WC, S Africa. Note: Under this name, all species with apparently paniculate inflorescences are kept together. A. lanceolatum has been attributed to Murray, who edited the 13th edition of Systema Vegetabilium . . . in 1774. Following Stafleu & Cowan (1981: 77), the author is Linnaeus, Murray being the editor only. A. papulosum  Galenia papulosa A. perfoliatum = Tetragonia decumbens A. procumbens = A. canariense

A. rigidum L.f. 1781: 261 T Thunberg 12056 (UPS!) = A. argenteum Ecklon & Zeyher 1837: 325  A. rigidum L.f. var. angustifolium Sonder in Harvey & Sonder 1862: 470 T Ecklon & Zeyher 2129 (B) = A. rigidum L.f. var. villosum Adamson 1959: 35 T Zeyher 2634 (S) = A. sericeum Fenzl ex Sonder in Harvey & Sonder 1862: 470 nomen nudum T Zeyher 2634c (?) Shrubs with smooth old stems, younger I with soft, appressed, long hairs; L alternate, elliptical, acuminate, narrowing to the base, covered densely with long, white hairs, 5–15 mm l, 3–8 mm b; Fl in cymes in the upper part of the branches, distant, perigon with spreading hairs outside, stamens numerous (Fig. 6); Fr sharply angled at the upper edges, 5 locules; Ecol in sand or gravel, mostly near the sea; Distr from Caledon, WC, to East London, EC, S Africa. Note: The varieties described differ mainly in the density of the hair cover, a feature variable in expression and changing during the aging of the leaves and stems. A. rigidum var. angustifolium = A. rigidum A. rigidum var. villosum = A. rigidum A. rodwayi  Gunniopsis rodwayi A. sarmentosum L.f. 1781: 260 T Sparrmann s.n. LINN 650.3 (LINN) = A. sarmentosum L.f. var. hirsutum Ecklon & Zeyher 1837: 327 T Ecklon & Zeyher 2140 γ (S) = A. sarmentosum L.f. var. strigosum Ecklon & Zeyher 1837: 327 T Ecklon & Zeyher 2140 ß (S). Shrubs with lax, procumbent branches 10–15 cm l, woody at their bases only, young I reddish-brown, with few to many white hairs, sometimes spreading; L opposite, cylindrical, connate at base, hair cover variable, 10–45 mm l; Fl in  terminal cymes, each with a pair of leaves below, perigon with white, appressed hairs outside, white or cream-coloured inside, cucullate, stamens in groups of 3–8; Fr

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3–6 mm b; Fl on short shoots, apparently solitary, perigon with scattered appressed hairs outside, glabrous inside, 3–6 mm l, stamens numerous; Fr obovoid, longer than wide, rounded at the top, 5 locules; Ecol on dry stony slopes or in river beds; Distr southern districts of Namibia. Note: Although occurring in a wide area, the species seems not to have been collected often. It is readily distinguished from all others by the long fruit.

Fig. 7 Aizoon schellenbergii with whitish erect to oblique branches ending in leafless tips with solitary capsules (Copyright H.E.K. Hartmann)

almost woody, turbinate, 4–7 mm ∅; Ecol in gravel or on more rocky ground, usually under bushes; Distr from Namaqualand, NC, to Cape Town and Swellendam, WC, S Africa. Note: The inclusion of Mesembryanthemum hexaphyllum Haw. 1821: 165 as a synonym of A. sarmentosum as supposed by Sonder (1862: 471) and Adamson (1959: 42) cannot be upheld. The drawing of Burman (1738: t. 26, f 2) is the cited type of the species, and that plant is definitely a member of the genus Portulaca. It should be noted that Sonder and Adamson cited above refer to different, wrong pages, and one also to a wrong year of publication. A. sarmentosum var. strigosum = A. sarmentosum A. sarmentosum var. hirsutum = A. sarmentosum A. schellenbergii Adamson 1959: 39  A. fruticosum Schellenberg 1913: 502 non L.f. 1781: 261 nomen illeg. T Dinter 887 (Z). Shrubs with erect to oblique branches, woody, stems 15–40 cm l, young I covered by short, white, appressed hairs, older I glabrous and whitish pale (Fig. 7); L alternate, obovate, narrowing into a short petiole, folded upwards, 5–15 mm l,

A. secundum  Galenia secunda A. sericeum = A. rigidum A. sessiliflorum = Aizoanthemum hispanicum A. spathulatum = A. canariense A. stellatum = A. paniculatum A. squamulosum  Galenia squamulosa A. teretifolium  Acrosanthes teretifolia A. theurkauffii = Mesembryanthemum cryptanthum A. tomentosum = A. paniculatum A. virgatum Welwitsch ex Oliver 1871: 584 T Welwitsch 1113 (K). Slender, erect shrubs with branches 50–75 cm l, I silky-tomentose, old ones with a white or grey bark, basally woody; L alternate, oblanceolate to obovate, covered by silvery, dense hairs, 15–50 mm l; Fl mostly as 3–5 in an axillary group, rarely solitary, 6 mm ∅, stamens 50–60; Fr obpyramidal, 4–5 locules; S 1–2 per locule; Ecol in dry rocky places, e.g. in river beds; Distr Benguela, Mossamedes, Angola; northern districts of Namibia. Note: Beside the erect, slender growth, the low number of seeds per locule is characteristic of the species. A. zeyheri Sonder 1862: 470 T Zeyher 717 (K!). Prostrate with ascending branches, stems woody, I with closely appressed, rather coarse white hairs; L opposite, sessile, ovate, covered by coarse, white, short hairs, 4–6 mm l, 2–3 mm b; Fl 2.25 mm l; Fr 2–3 capsules close together, similar to the fruits of A. canariense but hardly 5-angled, 5 locules, ∅ 2.5 mm; S 2 per locule, blackish; Ecol among rocks; Distr Namaqualand, WC, S Africa. Note: The species is morphologically similar to A. canariense, from which it can be distinguished

Aizoon AIZOOIDEAE

51

easily by the tiny white leaves, the tiny flowers, and the small fruit.

V. caulifloris = Aizoon canariense V. heisteri = Aizoon canariense

A. zygophylloides  Gunniopsis zygophylloides Veslingia  Aizoon

References

The name Veslingia Heister 1748: 11 has been created by the author in a list of plants with dry fruits, as a replacement “pro Ficoidea T. & Aizoon Linnaei”. Since this publication appeared before the starting point for nomenclatural regulations, Linnaeus’ Species Plantarum (1753), Veslingia Heister is not available as a name at genus level. Instead, the later publication of Fabricius contains the correct name: Veslingia Fabricius 1759: 201; the name should, therefore, be cited as Veslingia Heister ex Fabricius. The citations of Fabricius 363 by Willdenow (1799) and Moench (1802) refer to the second edition of the work: Fabricius 1763: 363. Fabricius did not cite any species under Veslingia Heister, but added immediately after the genus name: “Aizoon Linn.”, stressing the fact that both generic names are synonyms. The name Veslingia has been used again by Willdenow (1799: 1053), who listed “Veslingia Heisteri. Fabric. helmst. p. 363” as a synonym under Aizoon canariense, creating in this way perhaps unintentionally a new superfluous synonym. Moench (1802: 299–300) gave a description of the genus Veslingia and named one species: V. caulifloris Moench. Behind the description of this species, Moench cited the figures of Nissolle as “1711. icon. t. 19” (correctly: 1730: t. 13) and Volckamer “p. 236 icon. 236” (correctly: Volckamer 1718: t. 236), each showing a plant of Aizoon canariense. Furthermore, Moench (1802: 300) referred to Linnaeus’ description of Aizoon canariense (Linnaeus 1762: 1053), pointing out that Veslingia differed in the possession of free stamens and the number of stamens widely from the genus Aizoon. Of the cited figures, Nissolle. 1730: t. 13 was chosen as the lectotype, based on the fact that additional drawings of details permit a well-founded identification of the species (Hartmann 2001b: 349).

Adamson R.S. (1959) The South African species of Aizoaceae VII. Aizoon. JSAB 25: 29–51 Brown N.E. (1908) Diagnoses africanae XXIV. KB 1908: 290–291; XXV: 407 Burman J. (1738) Rariorum africanorum plantarum. . ., decas prima: 64, t. 26, f 1–3. Boussiere, Amsterdam Chinnock R.J. (1983) The Australian genus Gunniopsis Pax (Aizoaceae). Journal of the Adelaide Botanical Garden 6: 133–179 Christensen C. (1922) Index to Pehr Forsskål: Flora aegyptiaco-arabica 1775 with a revision of Herbarium Forskålii . . . Dansk Botanisk Arkiv 4, part 3: 1–54 Compton H. (1931) The flora of the Whitehill district. TRSSA 19: 283 Crantz H.I.N. (1766) Institutiones rei herbariae iuxta . . .1: 135–136. Kraus, Vienna Ecklon C.F., Zeyher C. (1837) Enumeratio plantarum africae australis extratropicae . . .3: Ficoideae: 308–329. The authors, Hamburg Fabricius P.C. (1759) Enumeratio methodica plantarum horti medici Helmstadiensis . . .: 201. Drimborn, Helmstedt Fabricius P.C. (1763) Enumeratio methodica plantarum horti medici Helmstadiensis . . . ed. 2: 363. Drimborn, Helmstedt Forskål P. (1775) Flora Aegytiaco-arabica sive descriptiones plantarum . . .: 25, 69–71, 95, CVIII, no. 200, t. 10. Möller, Kopenhagen Friedrich H.-C. (1960) Ficoidaceae. MBSM 3: 613–615 Friedrich H.-C. (1970) Aizoaceae. In: Merxm€ uller H. (ed) Prodromus einer Flora von S€ udwestafrika 27: 1–135 Gonçalves M.L. (1978) Aizoaceae sensu lato. Flora Zambesiaca 4: 508–555 Hartmann H.E.K. (2001a): Illustrated Handbook of Succulent Plants: Aizoaceae A-E. Springer. Heidelberg Hartmann H.E.K. (2001b): Illustrated Handbook of Succulent Plants: Aizoaceae F-Z. Springer. Heidelberg Harvey W.H., Sonder W. (1862) Flora Capensis 2: 386–478, 596–598, 616. Hodges, Smith & Co, Dublin Haworth A.H. (1795) Observations on the genus Mesembryanthemum . . ., part 2. The author, London Haworth A.H. (1803) Miscellanea naturalia, . . . J. Taylor, London Haworth A.H. (1821) Revisiones plantarum succulentarum. R. & A. Taylor, London Heister L. (1748) Systema plantarum . . .: 11. Drimborn, Helmstedt

52 Hill J. (1756) Aizoon. British Herbal: 53–54, f s.n. Osborne & Shipton, London Lamarck J.-B.P.A. (1792) Aizoon. Encyclopédie méthodique. Botanique . . . 3,2: 418. Panckoucke, Paris Linnaeus C. (1753) Species plantarum. Facs. 1957. Ray Society, London Lennaeus C. (1762–63) Species plantarum . . . (2nd ed.) Linnaeus C. (1774) In: Murray J.A. 1774: 392 Linnaeus C. fil. (1781) Supplementum plantarum. Systematis vegetabilium – ed. 13 generum plantarum: 227, 258–261, 392. At the expenses of the orphanage, Braunschweig Moench C. (1802) Supplementum ad methodum plantas . . .: 190–193, 299–300. Cotta, Marburg (reprint 1966: Koeltz, Königstein) Murray J.A. (1774) Caroli a Linné equitis Systema vegetabilium secundum classes . . . ed. 13: 388–392. Dieterich, Göttingen Nissolle G. (1730) E’Tablissement De Quelques Nouveaux Genres De Plantes. Histoire de l’Académie Royale des Sciences of the year 1711: 316–320, 324, t. 13, Paris

H.E.K. Hartmann Oliver D. (1871) Flora of tropical Africa 2: 582–588. Reeve & Co., London Petiver J. (1703/4): Gazophylacii naturae et artis decades X . . . 1: Decas IV/V: t. 39. Christophorus Bateman, London (copy seen from Göttingen) Schellenberg G. (1913) Aizoaceae africanae 2. BJS 48: 498–504 Schlechter R. (1897) Decades Plantarum Novarum AustroAfricanarum. Decas iv. JB 35: 280 Sonder O.W. (1862) Aizoon. Galenia. Mesembryaceae. Trianthema. In: Harvey W.H., Sonder W. (eds) Flora Capensis Sprengel C. (1827) Linnaei systema vegetabilium ed. 16, curae posteriores 4, pars 2: 195–196. Dieterich, Göttingen Stafleu F.A., Cowan R.S. (1981) Taxonomic literature 2nd ed. 3: 77, 670. Bohn, Scheltema & Holkema, Utrecht Volckamer J.G. (1718) Flora Noribergensis sive Catalogus Plantarum in agro norenbergensi . . .: 236, t. 236. Monath, N€ urnberg Willdenow C.L. (1799) Species plantarum, ed. 2: 1025–1053. G.C. Nauk, Berlin

Aloinopsis RUSCHIOIDEAE H. E. K. Hartmann

Aloinopsis Schwantes ZSK 1926: 177; Brown JB 1928: 77–78; Bullock 1939: 153–161; Bolus NM3 1958: 372–381; Hartmann 2001: 33–37; Hammer 2013: 12–53  Acaulon N.E.Br. JB 1928: 76 nomen illeg. non C.M€uller 1847: 99  Aistocaulon Poellnitz in Jacobsen 1935: 123 nomen illeg. Typus A. rosulata (Kensit) Schwantes  Acaulon rosulatum (Kensit) N.E.Br.  Aistocaulon rosulatum (Kensit) Poellnitz Etym Aloe, a monocotyledonous succulent with triangular, pointed leaves, Gk -opsis, aspect, appearance, resemblance to, in allusion to the leaf shapes of some of the originally included species, now considered members of the genus Nananthus. Compact with an irregularly thickened rootstock either fat and truncate or with some thin, short branches reaching to the surface, 2–3 leaf pairs to a branch in the resting state, forming a rosette; L with a rough surface from raised dots or with prominent warts above tannin idioblasts, flat, rarely trigonous with a flat, triangular area on top, calcium crystals thick in the epidermis of the warts or dots, thin to absent in the valleys in between, where the stomata are placed, wax in a  thick layer on the warts or dots, as horizontal plates in the valleys; Fl mostly appearing ebracteolate because the bracteoles are almost as

H.E.K. Hartmann (*) Biozentrum Klein Flottbek, Hamburg, Germany e-mail: [email protected]

long as leaves, K 5–6, petals vittate or not, then changing colour from base to tip, yellow, salmon, peach-coloured, pink, or cream, stamens in a broad cone over the stigmas, nectary as a ring; Fr with a rounded to sharply raised top and a funnel-shaped to almost semi-globose base, dispersed as tumble fruits with or without stalk and calyx lobes, the interior of the Titanopsis-type with very broad but short valve wings tapering into awns of the expanding keels, closing bodies absent or present, 8–12 locules; S maroon to dark brown, mostly almost round in side view; Ecol mostly on shales of the central S African plateau, rainfall expected mainly in March and November, in some species also in winter (Fig. 1); Distr part of WC, southern part of NC, western part of EC, S Africa. Note: Delineation and character set of the genus were under dispute over several decades (Brown JB 1928: 77–78, Bullock 1939: 153–161). Bolus (NM3 1958: 372–381) settled the uncertainties by a discussion of the relevant aspects giving also a list of the included species. She understood Aloinopsis in the wider sense (s.lat.) including the genera Prepodesma and Deilanthe, most probably based on similarities of outer capsule shape and locule numbers above five. More extensive studies of the complex indicate that numerous differences exist between the three taxa involved. Prepodesma and Deilanthe, both described by Brown (1930), are, therefore, treated here as separate genera leaving Aloinopsis s.str.

# Springer-Verlag GmbH Germany 2017 H.E.K. Hartmann (ed.), Illustrated Handbook of Succulent Plants: Aizoaceae, DOI 10.1007/978-3-662-49260-4_7

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H.E.K. Hartmann

Wh

Pi

February Lü

Jo Pc

Au

VR

March OR No

Ku

Up

Wa

Pm

VR

Gr

Ki

Summer March and November Winter

A.B.

Ag

P.N.

Po

Ke

Ma

OR

Sp

Ho

Ja

Ga

Bi

Va

Lo

OR Bu

Ca

Rc

Cl

L.B.

Su

To Wo

Ab

La

P.A.

Mo

C.T.

Ri Br

Ou Un Ge

Wi

B.E.

Mi

Um

Cr

G.R.

St

Gh

E.L.

Ui P.E.

All seasons

Fig. 1 Distribution of Aloinopsis in five different partial areas, each comprising distinct species, expecting rainfall in winter or in March and November (Copyright H.E.K. Hartmann).

Key to the Genera in Aloinopsis s.lat. 1. – 2.

–

Leaf surface rough from raised dots or warts, stamens in a cone, at least when opening, bracteoles apparently absent or inconspicuous Leaf surface velvety or smooth, not rough to the touch, stamens in an erect position from the beginning of the anthesis, bracteoles present Leaves velvety from long papillae, greyish from wax and crystals in the outer epidermal wall, apex and margins of leaves rounded, fruit with a thick columella in the centre, valve wings short, closing bodies present, 10–15 locules, seeds 1 mm long

with eight species only. A key is given to explain the differentiating character states of the three genera distinguished here. The distribution of Aloinopsis s.str. falls, according to new data, into four distinct separated parts, each with one or two species. In the latter case, the two species seem to be adapted to different soils and have never been found growing sympatrically. Aloinopsis s.str. can be divided after flower colours, as in the key below, or after leaf surfaces.

Aloinopsis s.str. 2 Deilanthe

Prepodesma

One group, consisting of A. acuta, A. loganii, A. rosulata, A. rubrolineata, and A. schoonesii, shows rough surfaces because elevations are formed above small groups of subhypodermal tannin idioblasts, giving a resistence to the touch. A second group comprises A. luckhoffii, A. malherbei, and A. spathulata, all exhibiting distinct white, hard warts, in which several tannin idioblasts are piled up forming a solid centre of each wart. When long, the protrusions can be bent, but not broken off easily.

Aloinopsis RUSCHIOIDEAE

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Key to the Species of Aloinopsis s.str. 1. – 2. – 3. – 4. – 5. – 6. – 7. –

Flowers vittate, i.e. with a central red stripe Flowers evittate, but changing colour from base to tip Leaves sunken in the ground, only the triangular tips visible Leaves at least partly visible above the ground Leaves  erect, tips rounded Leaves bent backwards completely, lying on the ground, tips pointed Leaves apically pointed Leaves apically rounded Leaves with conspicuous warts on the surface Leaves inconspicuously rough Leaves thick, thickness >1/2 width Leaves flat, thickness 300 mm rainfall p.a. in March and November; Distr Steytlerville, Willowmore, EC, Beaufort West, WC, S Africa. Note: Since the plants are easily overlooked in habitat, the distribution of the species is very incompletely known. Populations appear to occur widely dispersed, but are often rich in individuals. A. rubrolineata (N.E.Br.) Schwantes ZSK 1926: 178  M. rubrolineatum N.E.Br. KB 1911: 82  Nananthus rubrolineatus (N.E.Br.) Schwantes 1928: 68 T s.n. from Graaff-Reinet (K) = Nananthus cradockensis L.Bolus NM3 1939: 170 T James 1075/31 (BOL) = A. dyeri (L.Bolus) L. Bolus SAG 1929: 245  Nananthus dyeri L. Bolus NM2 1929: 83 T Dyer s.n. March 1929 (BOL) = A. jamesii L.Bolus NM2 1931: 294  Nananthus jamesii (L.Bolus) L.Bolus NM3 1938: 133 nomen illeg. non L. Bolus NM2 1931: 304 T James 1075/31 (BOL). Thick rootstock with up to 15 mm ∅; L rough from elevations above tannin idioblasts, widely recurved, the lower leaf surface touching the ground, triangular as seen from above, pointed, L c. 35 mm l, 17 mm b, 9 mm t; Fl ∅ 29–37 mm, K 5, 55–60 yellow petals with red stripes, 190–240 stamens; Fr the capsule base merging into a firm pedicel, base of capsule broadly funnel-shaped, the whole capsule

57

Fig. 2 Aloinopsis rosulata with two younger ripe fruits still with remains of their stalks and an old fruit from the previous year, demonstrating that the tumble fruits finally remain open, being able to loose their last seeds independently from rain (Copyright H.E.K. Hartmann)

including stalk dispersed as a tumble fruit, valve wings broad at base, narrowing into awns abruptly at c. 1/2 the length of the valve, closing body absent, 10–11 locules; S almost round in outline, dark brown, testa cells dome-shaped, S 1.15–1.25 mm l, 1 mm b; Ecol in silty flats or on shaly soils, c. 400 mm rainfall p.a. in March and November; Distr Graaff-Reinet, EC, S Africa. Note: A. rubrolineata is less susceptible to mould than other species, flowering regularly in cultivation in Europe. A. schooneesii L.Bolus NM2 1931: 295  Nananthus schooneesii(L.Bolus) L.Bolus NM3 1938: 134 T Schoonees 8657 = A. schooneesii L.Bolus var. acutipetala L.Bolus NM3 1958: 381 T van Blerk 2444/36 (BOL) = A. schooneesii L.Bolus var. willowmorensis L.Bolus NM3 1958: 381 T Chipp 1313/32 (BOL). The very thick rootstock developing several fragile stems reaching up to the surface; L trigonous with a marked chin, the apical area flat and level with the soil surface, elevations above tannin idioblasts with crystals in the outer epidermal walls, L c. 17 mm l, 7 mm b, 4 mm t; Fl with bracteoles c. as long as the leaves, K 5, petals yellow with a red stripe; Fr details not known; Ecol silty flats, c. 200 mm rainfall p. a. in March and November; Distr Willowmore, EC, S Africa. Note: In the group of species sharing a rough leaf surface without proper warts, A. schooneesii

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is the smallest with the relatively thickest leaf, growing sunken deeper into the ground than the others. In leaf shape, it resembles most closely A. luckhoffii, which is, however, bigger and possesses warts. A. schooneesii var. acutipetala = A. schooneesii A. schooneesii var. willowmorensis = A. schooneesii A. setifera = A. luckhoffii A. spathulata (Thunberg) L.Bolus NM3 1954: 256  M. spathulatum Thunberg 1791: 5  Titanopsis spathulata (Thunberg) Schwantes ZSK 1927: 105  Nananthus spathulatus (Thunberg) G.D.Rowley NCSJ 1978: 6 LT, designated by Brown, Gardeners’ Chronicle 1931: 14, Thunberg s.n. (UPS!) = A. crassipes (Marloth) L.Bolus SAG 1929: 245  M. crassipes Marloth FPSA 1922: t. 73  Titanopsis crassipes (Marloth) N.E.Br. GC 1931: 14  Nananthus crassipes (Marloth) L.Bolus NM3 1938: 133 T Marloth 9681 (?). The thick rootstock widening to the top, splitting into several branches reaching the surface; L erect, flat, warts moderately developed, papillae restricted to elevations, wax cover breaking into plates on the elevations, rugose-continuous in the valleys, many platelets, L 10–20 mm l, 8–10 mm b, 2.5–3 mm t; Fl ebracteolate, 25–30 mm ∅, K 5, petals pink or white (?); Fr top raised abruptly in the centre, the tips of the valves forming awns, the base semi-globose, the stalk breaking off early from the nearly semi-globose base (Fig. 3), columella thick, valve wings narrowing into awns distally, closing body absent, but a bulge in its place, 10–11 locules; S short pear-shaped to triangular in outline, medium to dark brown, testa cells elevated to dome-shaped, S 0.9–1 mm l, c. 0.7 mm b; Chr 2n = 18; Ecol in shaly flats, c. 125 mm rainfall p.a. in winter or in March and November; Distr Sutherland, NC, S Africa (Fig. 3). Note: In the moderate expression of the warts similar to A. villetii, but differing clearly in the flat leaves. Under darker and wetter conditions than in habitat, five and more leaf-pairs per season can be formed, but flowers are rare.

Fig. 3 Aloinopsis spathulata with one of three capsules still on its stalk, rhs, the other two broken off their stalks and turned over as a tumble fruit (Copyright H.E.K. Hartmann)

A. thudichumii  Deilanthe thudichumii A. villetii = A. luckhoffii A. wilmaniae = Nananthus aloides Acaulon  Aloinopsis Brown (JB 1928: 76–77) established this name in order to replace Aloinopsis Schwantes because “Schwantes’ name cannot be maintained on account of the entirely false characters he assigned to it”. However, since Brown included as the only species the type species of Aloinopsis, A. rosulatum, the new name is illegitimate (McNeill et al. 2012: Art. 52.2). Furthermore, the name Acaulon C. M€uller (1847: 99) had been published earlier for a moss; Acaulon N.E. Br. is therefore illegitimate under Art. 53.1 (McNeill et al. 2012), too (Hartmann 2001: 23). Acaulon rosulatum  Aloinopsis rosulatum Aistocaulon  Aloinopsis Poellnitz (1935: 123) established this genus name as a replacement of the illegitimate name Acaulon N.E.Br. because that name is a homonym of Acaulon C.M€uller, a moss genus (McNeill et al. 2012: Art. 53.1). Both names, Acaulon N.E. Br. and Aistocaulon Poellnitz, are also illegitimate names for the older name Aloinopsis since all three are based on the same type species (McNeill et al. 2012: Art. 52.2) (Hartmann 2001: 27). Aistocaulon rosulatum  Aloinopsis rosulatum

Aloinopsis RUSCHIOIDEAE

References Bolus H., Kensit L. (1909) Contributions to the African flora. TRSSA 1: 147–163, t. 21 Bolus L. (1928–1935) Notes on Mesembrianthemum and allied genera. NM2: 1–508. 1–16 (9.11.1928), 17–32 (21.12.1928), 33–48 (24.1.1929), 49–64 (12.4.1929), 65–80 (3.5.1929), 81–94 (6.6.1929), 95–110 (4.7.1929), 111–129 (16.8.1929), 130–146 (4.11.1929), 147–160 (22.11.1929: the last two texts end in the middle of a description), 161–176 (20.2.1930), 177–192 (9.5.1930), 193–208 (15.8.1930), 209–224 (12.11.1930), 225–244 (12.2.1931), 245–268 (1.5.1931), 269–292 (3.7.1931), 293–308 (24.9.1931), 309–336 (29.1.1932), 337–356 (24.6.1932), 357–376 (5.12.1932), 377–396 (19.5.1933), 397–416 (16.10.1933), 417–436 (26.1.1934), 437–452 (23.5.1934), 453–472 (17.8.1934), 473–508 (11.2.1935). 517–520 Corrigenda. University of Cape Town. Bolus Herbarium. Cape Town Bolus L. (1929) Mesembrianthemum and allied genera. SAG 19: 49, 181, Lithops fulleri var. Tapscottii: 182. (figure and legend only), 244–245, 288 Bolus L. (1936–1958) Notes on Mesembryanthemum and allied genera. NM3: 1–19 (5.2.1936), 21–44 (31.7.1936), 45–78 (14.1.1937), 79–114 (12.8.1937), 115–138 (24.3.1938), 139–188 (20.7.1939), 189–236 (25.8.1950), 237–288 (30.8.1954), 289–417 (30.4.1958). University of Cape Town. Cape Town Bolus L. (1947) Ruschia maxima. FPSA 26: t. 1013, Delosperma jansei 26: t. 1024, Aloinopsis malherbei. FPSA 26: t. 1035 Brown N.E. (1911) Diagnoses africanae XXXIX, XLV. KB 1911: 82–83, 313, 356–359 Brown N.E. (1928) Mesembryanthemum and allied genera. JB 66: 75–80, 106–107, 138–145, 171–172, 267–268, 322–327 Brown N.E. (1930) Mesembryanthemum . . . GC 87: 13–14, 32–33, 71–72, 126, 151, 186, 211–212, 267, 515–516; GC 88: 8, 278–280, 474, 513, 516

59 Brown N.E. (1931) Mesembryanthemum . . . GC 89: 13–15, 53–54, 92–93, 133–135, 259, 279, 294, 389; GC 90: 13–14, 136–137, 335 Bullock A.A. (1939) Notes on Mesembryanthemaceae: II. The lectotypes of Nananthus and Aloinopsis. KB 52: 153–161 Hammer S. (2013): Mesembs. The Titanopsis Group. Little Sphaeroid Press. Oakland, California Hartmann H.E.K. (2001): Illustrated Handbook of Succulent Plants: Aizoaceae A-E. Springer. Heidelberg Jacobsen H. (1935) Succulent plants. Williams & Norgate, London Kensit L. (1909) In: Bolus H., Kensit L. (1909) Marloth R. (1922) Mesembryanthemum crassipes. FPSA t. 73 McNeill J., Barrie F.R., Buck W.R., Demoulin V., Greuter W., Hawksworth D.L. Herendeen P.S., Knapp S., Marhold K., Prado J., Prud’homme van Reine W.F., Smith, G.F., Wiersema J.H. et al. (2012) ICN International Code of Nomenclature for algae, fungi and plants (Melbourne Code). Regnum Vegetabile 154. A.R.G. Gantner Verlag. M€ uller C. (1847) Systema muscorum ordinis “Cleistocarpi” Brid. Botanische Zeitung 5: 97–102 Poellnitz K. von (1935) In: Jacobsen H. (1935) Rowley G.D. (1978) Reunion of some genera of Mesembryanthemaceae. NCSJ 33: 6–9; A postscript: 62 Schwantes G. (1926) Zur Systematik der Mesembrianthemen. ZSK 2: 173–189, 242, 277 Schwantes G. (1927) Zur Systematik der Mesembrianthemen. ZSK 3: 14–30, 105–106 Schwantes G. (1928) Mesembriaceen unserer Kulturen in neuer Benennung. GF 77: 68–69; Chasmatophyllum nelii 43–44; Nelia meyeri spec. nov.: 129–130 Thunberg P. (1791) Descriptiones Mesembryanthemorum quorundam in Capitis Bonae Spei Africes . . . Novorum Actorum medico-physicorum academiae Caesareae Leopoldino-Carolinae naturae curiosorum in Germania 8: App.: 1–18

Amphibolia RUSCHIOIDEAE H. E. K. Hartmann

Amphibolia L.Bolus ex Herre 1971: 70; L.Bolus JSAB 1965: 169 nomen invalid.; Toelken & Jessop 1976: 64; Hartmann & Dehn 1989: 179–182; Hartmann Bradleya 1996: 29–56; Hartmann Bradleya 1998: 44–51; Hartmann 2001: 37–39 Lectotypus A. maritima L.Bolus ex Toelken & Jessop = A. laevis (Aiton) H.E.K.Hartmann Etym Gk amphibolia, uncertainty or doubt, because the combination of closing bodies and broad valve wings occurring also in Stoeberia raised doubts about the placement of the type species. Low shrubs to c. 40 cm tall, ∅ to 70 cm, or higher and not so broad, I white to yellowish or greyish; L club-shaped to trigonous, epidermis smooth, stomata hardly sunken, outer walls rather thin, with a thin layer of calcium oxalate sand, often grey from wax; Fl 1–3 per cyme, bracteate and bracteolate, petals pink with a complicated marking: the basal darker line lies in a lighter whitish or greenish rim separating the line from the outer dark purple area, in some species petals uniformly pink or white, filamentous staminodes often apically dark purple, surrounding the central stamens in a distinct, broken cone resembling a pitchershape in fresh flowers: the bases of the filaments almost horizontally arranged, the upper halves

H.E.K. Hartmann (*) Biozentrum Klein Flottbek, Hamburg, Germany e-mail: [email protected]

turning upwards rather abruptly near the middle, stigmas short, densely papillate outside as well (Fig. 2), nectary in a lophomorphic ring; Fr always persisting on the plant, base funnelshaped, merging into the stalk, valve wings very broad and rectangular, covering membranes complete with a low distal recurving under which closing rodlets or closing ledges are placed, closing bodies small, 5, rarely 6 locules; S glabrous and almost round; Ecol in sand, gravel, or rocks, from below 100 to above 300 mm rainfall p.a. in winter, fog from the sea (Fig. 1); Distr L€uderitzSouth, Namibia; districts along the W coast of S Africa. Note: Hartmann (1993: 63) placed Amphibolia as a synonym with Eberlanzia, a view not upheld here. The less xeromorphic leaves, the peculiar construction of the androecium and the stigmas being papillate on their outside, the very broad valve wings and the funnel shaped base of the fruit of Amphibolia permit at present a clear distinction and circumscription. Features shared with Eberlanzia are the whitish stems and the glabrous, almost round seeds. Amphibolia should be studied again in its relation to the genera Stoeberia (also with very broad valve wings) and Ruschianthemum, both with whitish stems as well. The complex of the four genera, until now included in the Ruschia-group (Hartmann 1991, 1993), may represent an evolutionary branch of its own. Amphibolia and the

# Springer-Verlag GmbH Germany 2017 H.E.K. Hartmann (ed.), Illustrated Handbook of Succulent Plants: Aizoaceae, DOI 10.1007/978-3-662-49260-4_8

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Wh

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OR No

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Fig. 1 Distribution of Amphibolia is restricted to coastal or coast-near habitats along the south-western coast of southern Africa, where saline spray and morning fog from the sea can be expected regularly (Copyright H.E.K. Hartmann) Key to the Species of Amphibolia 1. – 2. – 3. – 4. –

Capsules with 5 locules, 6 occurring only as an exception in few populations, base of fruit long, funnel-shaped Capsules with 6 locules throughout, base of fruit short, funnel- or bell-shaped Leaves short (length at most double the breadth or width), trigonous, densely set, base of fruit mostly enclosed by bracteoles Leaves more than twice, mostly 3–4 times as long as broad or thick, club-shaped, base of fruit raised well above the bracteoles Leaves slender, 15–30 mm long, 4–5 mm broad and thick Leaves thick, 10–20 mm long, 8–10 mm thick, often crescent-shaped in side view Capsule with a distinct closing rodlet at the distal end of the covering membrane, closing body hook-shaped Capsule with a closing ledge at the distal end of the covering membrane, tiny closing body knobshaped

supposedly related species need a revision to clarify the taxonomy with confidence. Most species grow allopatrically, but A. rupis-arcuatae and A. succulenta are in some places found together. A. gydouwensis  Phiambolia incumbens A. hallii  Phiambolia hallii A. hutchinsonii = A. laevis

2 A. obscura A. rupisarcuatae 3 A. succulenta 4 A. laevis A. saginata

A. laevis (Aiton) H.E.K.Hartmann  Mesembryanthemum laeve Aiton 1789: 187 nec Thunberg 1791: 16 nomen illeg. NT, designated by Hartmann 2001: 38, drawing 191/142 M by Duncanson 31.6.1825 (K!) = Ruschia hutchinsonii L.Bolus NM2 1928: 23  M. hutchinsonii (L.Bolus) N.E. Br. GC 1930: 32  A. hutchinsonii (L.Bolus) H.E. K.Hartmann Bradleya 1998: 46 T Hutchinson

Amphibolia RUSCHIOIDEAE

63

placed in the genus Amphibolia. References to this species include: Gmelin 1791: 845; Haworth 1795: 254–255; Willdenow 1799: 1044; Haworth 1803: 64–65; Aiton 1811: 232–233; Haworth 1812: 302–303; Haworth 1821: 154–155. Of these authors, Willdenow indicated a purple flower, followed by Haworth 1795 and Aiton 1811. Ruschia promontorii is placed here in synonymy based on similarities in leaf surface only, because fruits are absent and the flowers are inadequately preserved. A. laevis is very similar to A. saginata and A. succulenta, all three perhaps subspecies of one species.

Fig. 2 Amphibolia laevis with an urceolate flower and the typical colouring (Copyright H.E.K. Hartmann)

288 (BOL!) = A. maritima L.Bolus ex Toelken & Jessop 1976: 64; L.Bolus 1965: 169 nomen invalid.  Ruschia maritima (L.Bolus ex Toelken & Jessop) G.D.Rowley NCSJ 1978: 62 T Hall 2885 (BOL!) = Ruschia promontorii L.Bolus NM2 1929: 121  M. pansifolium N.E.Br. GC 1930: 32 T Rohland 1543/20 (BOL!). Low shrubs to 20, rarely to 40 cm h, ∅ to 70 cm, adventitious roots on decumbent branches, short side branches erect, I whitish to yellowish; L light green to yellowish grey, club-shaped, emarcescent old ones remaining on the stem for some time, 10–20 mm l, 6–8 mm b, 6–10 mm t; Fl petals pale pink (Fig. 2); Fr base long, funnelshaped, closing rodlets prominent at the end of the covering membranes, closing body hook-shaped, 5 locules; S 0.9–1 mm l, c. 7.5 mm b; Ecol gravelly coastal plains, up to 350 mm rainfall p.a. in winter, fog; Distr from Vredendal to Simonstown, WC, S Africa. Note: Although it has been known since 200 years that the name M. laeve had been used for two different species, it was only a decade ago found that a drawing agreeing with the description of Aiton is kept at Kew. Based on the club-shaped leaves withering slowly, the figure is chosen as the neotype of the species which can, therefore, been

A. littlewoodii  Lampranthus mutatus A. maritima = A. laevis A. obscura H.E.K.Hartmann Bradleya 1998: 47 T Hartmann et al. 20805 (HBG!). Broad shrubs to 40 cm h, ∅ to 70 cm, I 35–40 mm l, ∅ to 5 mm; L bright light green, club-shaped to terete, 15–45 mm l, 5–10 mm b and t; Fl 1–3, ∅ 18 mm, bracteoles hooded, embracing the base of the flower, K 6, petals purplish-pink, 8 mm l, to 1.5 mm b, filamentous staminodes absent (?), anthers white; Fr base short, funnel-shaped, top raised, but rims low, covering membranes with thick and long closing ledges, closing body nearly hook-shaped, 6 locules; S 1.1 mm l, 0.8 mm b; Ecol on rocky slopes in shallow places,